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JOURNAL
OF
SOUTH AFRICAN
BOTANY
VOLUME 44 1978
Published under the authority
of the
Trustees
of the
National Botanic Gardens
of
South Africa,
Kirstenbosch,
Newlands, C.P.
Editor:
Prof. H. B. Rycroft
Director,
National Botanic Gardens
of
South Africa,
Harold Pearson
Professor of Botany
University
of
Cape Town
Publication dates:
44 (1): 1-— 96 [6th January, 1978]
44 (2): 97-202 [18th April, 1978]
44 (3): 203-315 [25th July, 1978]
44 (4): 317-421 [30th October, 1978]
JOURNAL
OF
SOUTH AFRICAN BOTANY
VOLUME 44 1978
CONTENTS
A CHECKLIST OF THE PLANTS OF NDUMU
GAME RESERVE, NORTH-EASTERN
TAUIUUTL/ANNID), [E; Sh POGUE cca acossseccccaaa
HABITAT OF THREATENED PROTEACEAE
ENDEMIC TO WESTERN CAPE COASTAL
PL/I. Uy Wo INTIS coo ccccccccncon0n0s
STUDIES IN THE GENUS CASS/A IN SOUTH
AFRICA: 2. NOTES ON CASSIA ITALICA
(MILL.) LAM. EX F. W. ANDR. K. D.
Gordon=Grayietie wee oceans
NUCLEOLAR HETEROCHROMATIN IN
ENCEPHALARTOS. D. J. Mogford.........
TWO NEW SPECIES AND A NEW COMBI-
NATION IN THE GENUS KALANCHOE. H.
PTOI KO Micke cee testes ah ear lesser keh meteae
BOOK REVIEWS: Glossary of genetics and
cytogenetics, classical and molecular, by R.
Rieger, A. Michaelis and M. M. Green (J. S.
Allan). Kryptogamen, by K. Esser (K. T. van
Wiaitmelo) ie 25 a Aestett cecsru teat wrecsrnaden acters
THE REPRODUCTIVE POTENTIAL OF
ERICA JUNONIA BOLUS: MEGA-
SPOROGENESIS AND MEGAGAMETO-
GENESIS. B. L. Robertson and J. E.
MceNalightOnipw ter etcantnna ce 02, nacre
EFFECT OF VARIOUS TREATMENTS ON
GERMINATION OF DORMANT SEEDS OF
STRELITZIA REGINAE AIT. H. A. van de
WGI AG RRR nS RELAIS Syrtins Eee scien RIE oee. cee
CENTROMERIC HETEROCHROMATIN IN
LACHENALIA TRICOLOR (L.) THUNB. D. J.
Mogford ssa tanh aaceete eae an ae ener abe
THE PECULIAR LEAF TRICHOMES OF
PETREA VOLUBILIS (VERBENACEAE). C.
Puff
LEAF ANATOMY OF JUBAEOPSIS CAFFRA
BECCR Be RODCiLSOMmee eee
A PRELIMINARY STUDY OF THE NITROGEN
NUTRITIONAL STATUS OF MEMBERS OF
THE SOUTH AFRICAN PROTEACEAE. O.
A. M. Lewis and W. D. Stock..............
A NEW SPECIES OF PECTINARIA. P. V.
Bruyns
Page
55
67
83
89
93
Sil
103
111
119
SOME PROBLEMS OF SOUTH AFRICAN
PTERIDOPHYTA. W. B. G. Jacobsen......
“DRIFT SEEDS” AND “DEAR DIRTY DUB-
LIN’: CORRESPONDENCE BETWEEN
JOHN MUIR AND MATILDA C. KNOWLES:
11930 —19Sou = CaINeISOnie ee
BOOK REVIEWS: Pollination mechanisms,
reproduction and plant breeding, by R.
Frankel and E. Galun (H. A. van Niekerk).
Proceedings of the Second National Weeds
Conference of South Africa (M. J. Wells) ...
THE GENUS GALIUM L. (RUBIACEAE) IN
SOUTHERN AFRICA. C. Puff.............
PHYTOSOCIOLOGICAL STUDIES ON TABLE
MOUNTAIN, SOUTH AFRICA: 1. THE BACK
TABLE. J. Glyphis, E. J. Moll and B. M.
CGampbellies ances oeincetenns ocho:
PHYTOSOCIOLOGICAL STUDIES ON TABLE
MOUNTAIN, SOUTH AFRICA: 2. THE
FRONT TABLE. D. Laidler, E. J. Moll, B. M.
Gampbelliand JNGlyphis...2.q.0s-e2 sae -
AN INVESTIGATION INTO THE ECOLOGY
AND PRODUCTIVITY OF EPIPHYTIC
MOSSES. N. H. G. Jacobsen .............
BOOK REVIEWS: Dormancy and the survival
of plants, by T. A. Villiers (J. A. de Bruyn).
Compositae in Natal, by O. M. Hilliard (B.
Nordenstam). Wood structure in biological
and technological research, edited by P.
Baas, A. J. Bolton and D. M. Catling (A. R. A.
NOG I) estar eevee eee scr sreintore Crewe nen
PRELIMINARY ANNOUNCEMENT: Thirteenth
International Botanical Congress ..........
THE VASCULAR ANATOMY OF THE SEED-
LING OF ENCEPHALARTOS EUGENE-
MARAISII VERDOORN. P. J. Robbertse, S.
van der Westhuizen and G. K. Theron .....
STUDIES IN THE GENERA OF THE D/OS-
MEAE (RUTACEAE): 7. |. Williams ........
HABITAT OF THREATENED SPECIES OF
SERRURIA AND PROTEA ENDEMIC TO
WESTERN CAPE COASTAL FLATS. A. V.
RATE US le os re casa ciane sero esiioresencnen
FURTHER NOTES ON PROTEA IN SOUTH
AERIGAMUSEAROUIKEG): 6 ccc nc ons momen:
FLAVANOIDS OF THE SUMMER RAINFALL
PROTEAS AND THEIR CHEMOTAXO-
NOMIC SIGNIFICANCE. C. W. Glennie
atq(o| (Le [E. IDEMPBOMN sscnconcssonavon00Dce
CHROMOSOME NUMBERS IN LACHENALIA
(LILIACEAE). R. Ornduff and P. J. Watters .
Page
157
187
201
203
281
291
297
313
315
317
329
363
373
381
387
TEN NEW SPECIES OF LACHENALIA
(LILIACEAE). W. RB. Barkeh cirri:
THE IDENTITY OF POLYPODIUM POP-
PIGIANUM METT. (FILICES) J. P. Rourke
and’ E. A; C.E.JEsSchelpe ssn 7 aeeeree.:
BOOK REVIEW: Mesquite: its biology in two
desert ecosystems, edited by B. B. Simpson
(Gs Vi Joubert): . sce a as Saar ee
INDEX TO PLANT NAMES: Vol. 44 (1-4) ....
Indexed in Biological Abstracts and Current
Advances in Plant Science.
391
419
421
423
THIS VOLUME IS DEDICATED TO
JOHN PATRICK MICKLETHWAIT BRENAN (1917— )
MA. B.Sc, LS. FIL Biol:
(Director of the Royal Botanic Gardens, Kew)
a specialist and authority on African plants, especially the families Commelinaceae,
Steruliaceae, Chenopodiaceae and Leguminosae; he has made extensive observations
and collections throughout Tropical and Southern Africa which form the bases for
numerous publications to his credit, these include a descriptive check list of the Woody
Plants of Tanganyika (1949) published in collaboration with P. J. Greenway as well as
valuable contributions to the Floras of Tropical East-Africa, West Tropical Africa and
Flora Zambesiaca; he has been associated with a number of important scientific
societies, having been Botanical Secretary of the Linnean Society of London from
1965-72, President of the Association of Tropical Biology in 1970—71 and is also a
member of the Library and Scientific Committees of the Royal Horticultural Society;
throughout his botanical career he has kept a close liaison with Kirstenbosch and other
major South African Botanical Institutes, to which he has given freely of his advice
concerning matters of policy and management.
Digitized by the Internet Archive
in 2016 with funding from
South African National Biodiversity Institute Libraries
https://archive.org/details/journalofsouthaf44unse
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FIGURE 2
Map of the vegetation of Ndumu Game Reserve
Simplified from the ‘Map of the vegetation formations of Ndumu Game Reserve’
by De Moor et al (which was drawn directly from the 1964 aerial photographs)
Esk 1977 |
SCALE 1: 36,000 APPROXIMATELY
JIS. Afr. Bot. 44 (1): 1-54 (1978)
A CHECKLIST OF THE PLANTS OF NDUMU GAME RESERVE,
NORTH-EASTERN ZULULAND
E. S. POOLEY
(St. Lucia Estuary, Natal)
ABSTRACT
An annotated checklist of the plants of Ndumu Game Reserve, northern Zululand is
given together with a brief floristic analysis and a description of the vegetation types of the
area. A map is included.
UITTREKSEL
"N LYS VAN DIE PLANTE IN DIE NDUMU WILDRESERVAAT IN NOORD-OOS
ZOELOELAND
‘n Lys van plante met aantekeninge in die Ndumu Wildreservaat, Noord Zoeloeland,
word aangebied met ’n kort floristiese ontleding en ’n beskrywing van die plantegroei tipes
van die area. ’n Kaart word ingesluit.
INTRODUCTION
The material for the checklist was collected during the period 1967 to 1974.
The specimens have been routinely named in the Department of Botany, Univer-
sity of Natal, Pietermaritzburg and are housed in that herbarium, with many
duplicates in the Natal Parks Board Herbarium, Hluhluwe Game Reserve. Other
collections cited are also housed in these two herbaria, namely those of E. J. Moll,
T. B. Oatley, J. M. Ross, K. L. Tinley, C. J. Ward and some others.
The plants in the checklist are limited to those of the game reserve in spite of
the fact that some most interesting material has also been collected just to the west
on the Lebombo foothills, and from the Sand Forest areas just to the east.
GENERAL DESCRIPTION OF GAME RESERVE
Ndumu Game Reserve is 10 100 ha in extent and is situated in north-eastern
Zululand (Tongaland) on the border between South Africa and Mozambique (Fig.
1). It lies at the southernmost extension of the Mozambique plain at approximately
latitude § 26° 53’ longitude E 32° 15’, about 70 km from the sea.
The Usutu river, known as the Rio Maputo in Mozambique, serves as part of
the international border as well as the game reserve boundary on the north. The
game reserve is fenced on the south, east and west. The Pongola river flows
through the reserve from the south boundary to its new confluence with the Usutu
Accepted for publication 27th June, 1977.
1
2 Journal of South African Botany
Durban
Scale 1 :4,000,000
Figure 1
Location of Ndumu Game Reserve
Checklist of Plants in Ndumu Game Reserve 3
in the north. The old course of the Pongola serves as a flood water feeder channel
to Inyamiti, iHotwe and Sabatane pans, either directly from the Pongola or as
feed-back from the Usutu river, depending on water levels in the two rivers. There
are several semi-permanent pans; Fontana, Shokwe, Bantana, Mvutsheni and
Banzi on the Usutu; and isiHilibindini, Mpumpununa, iPolwe, Inyamiti, iHotwe,
Sabatane, Ndwanini and Bakabaka on the Pongola, which are filled by these
perennial rivers. The extensive floodplains form approximately a third of the
reserve.
Its highest point is Ndumu Hill which rises from the Pongola floodplain (25 m
above sea level) to a height of 160 m above sea level. The foothills of the
Lebombo rise in the extreme west of the reserve. There are flats in between these
two elevated areas, with extensive areas of tall thicket with scattered trees, low
thicket with few trees (termed “‘scrub’’ in checklist), woodland and savanna.
The game reserve was proclaimed in 1924. Until 1959, about 1 000 Africans
and their goats and cattle lived there. Crops were cultivated mainly on the
floodplains. During 1959, all stock were removed and subsequent recuperation of
the vegetation was phenomenal (Tinley, 1964). The last Africans were moved out
of the reserve in 1966, but there is still considerable evidence of old kraal sites
and, especially on the Usutu Floodplain, of disturbed vegetation in old fields.
Arson has always been a factor in the life of the game reserve. Fires are put in
by poachers with regularity, particularly in the west but also, on occasions, on
Ndumu Hill.
The climate is sub-tropical, with a summer rainy season lasting from October
to April, although onset, duration and amount are very variable. In winter, dense
morning mists occur as a fairly regular feature. Mean monthly rainfall and
temperature data are contained in Table 1 (from De Moor et al., in press).
TABLE 1.
Average monthly rainfall and temperature data from Ndumu Game Reserve
=. ee ee
Rainfall (mm) Temperature (° C)
Month (1956-1971) (Max. + Min. 1968-1971)
2
NAWUAL Veen econ. cee 3. 2 44,4 27,1
ebruiany 4 oer ee 79,7 26,8
IWET(G) uben cteseeine ot che tie eee 61,2 25,8
INDE Meh tee A ts imc: 9 43,4 23,8
Wiciy ier ere oe cise ere cess oe = 17,2 20,4
Fre See See aa eee 14,7 18,1
tik ea Pe cert e282. 10,7 19,1
/-VEO CHE 22 2 OO eee Oe 14,4 20,6
Septenibeme cote. hs: 23,4 22,2
October Se 74,4 24,3
Naveniber aetiersacici ae2s 75,3 24,3
Decemiberan vaste oe cers 75,9
TOTAL AVERAGE ANNUAL RAINFALL: 534,7 mm
4 Journal of South African Botany
VEGETATION
A vegetation and soil survey was conducted in 1971 and 1972 by De Moor and
his collaborators, and twenty-nine vegetation formations were described according
to physiognomic characteristics (following Fosberg, 1967). A vegetation and soil
map is included in that work. For the purposes of this paper the game reserve will
be described following the main eight formations delimited in the De Moor
survey, but in less detail and drawing attention to areas of particular botanical
interest. A map is included showing the Zulu names for the various parts of the
game reserve and showing the main vegetation formations (Fig. 2).
Riverine Forest
This occurs as a broken strip of forest along the banks of the Usutu and
Pongola rivers, typically found on levées formed by floodwaters. The main tree
species are Ficus sycamorus, Rauvolfia caffra, Trichilia emetica, Syzygium
guineense, Adina microcephala, Ekebergia capensis, Sapium ellipticum and Phyl-
lanthus flacourtioides, which form a discontinuous canopy. The sub-canopy can
include Antidesma venosum, Phoenix reclinata, Kraussia floribunda, Tabernae-
montana elegans, Oncoba spinosa and Gardenia cornuta. Shrubs and climbers
include Allophyllus decipiens, Grewia caffra, Ficus capraefolia, Acacia
schweinfurthii, Monanthotaxis caffra, Pisonia aculeata, Capparis tomentosa, Ipo-
moea digitata, and Gymnema sylvestre. Herbs and grasses found are Justicia
glabra, Achyranthes aspera, Setaria chevalieri and Oplismenus hirtellus.
The area of forest described in De Moor et al., with a continuous canopy of 30
m, is the only such undisturbed area of forest of any size—and is now claimed by
Mozambique as it is in the confluence area of the Usutu and Pongola rivers and is
cut off by two of the old courses of the Pongola rivers.
The new course of the Pongola river has no Riverine Forest, but forest species
from the old course are spreading steadily down the new course.
Usutu Floodplain
Much of this floodplain is not inundated regularly, only at times of exception-
ally high flood levels. The composition of grass and sedge species differs greatly
from the Pongola Floodplain and is much more diverse. There are many channels
which are flooded most seasons; a noteworthy one being the Mtikeni which links
the Usutu river to the Mdipini pools, Bantana, Mvutsheni and Banzi pans. These
channels, being regularly flooded, support mostly wetland species which include:
Phragmites spp., Cyperus spp., Pennisetum glaucocladum, Sorghum verticilliflo-
rum, Hemarthria altissima and Typha latifolia. Patches of Riverine Forest are
found across the floodplain, and single large trees of the species Acacia albida, A.
tortilis, Sclerocarya caffra, Bolusanthus speciosus, Kigelia africana, Loncho-
carpus capassa, Ficus sycamorus and Trichilia emetica occur. The common
grasses on the sandy alluvium are Cymbopogon validus, Hyperthelia dissoluta,
Checklist of Plants in Ndumu Game Reserve 5
Themeda triandra, Aristida congesta subspp. congesta and barbicollis, Sporo-
bolus fimbriatus, Eragrostis superba, Trichoneura monachne and Cynodon dacty-
lon, with shrubby thickets of Pluchea dioscoridis, Vernonia colorata, Lippia
javanica and Securinega virosa.
Pongola Floodplain
This floodplain before it was affected by Jozini dam was either inundated for
several weeks each summer or sometimes not at all. However, once the dam
started controlling the flow of the river the floodplain was inundated to its highest
levels at unseasonal times and for extended periods of time. This has had a great
effect on the composition of grasses and sedges, which are changing steadily. On
wet mud (and over water) Echinochloa pyramidalis (prostrate form), Hemarthria
altissima and Cyperus fastigiatus predominate with the herb Grangea maderaspa-
tensis. The grasses along the edge of the floodplain and on raised levées are
mostly Eriochloa meyerianum (= Panicum meyerianum), Panicum coloratum,
Chloris gayana, Sorghum verticilliflorum, Paspalum commersonii, Eriochloa
stapfiana, Urochloa mossambicensis, Digitaria spp., Echinochloa pyramidalis (erect
form). This area is heavily utilised by hippo, and in winter, when grazing
elsewhere is low, rhino, nyala, impala and reedbuck can be found in large
numbers. Sesbania sesban, Phragmites spp., and Ficus capraefolia form a discon-
tinuous transitional area between forest and grasses. East of the Pongola near
Mavilo, spurwing geese can be found in great numbers in the shelter of the
reedbeds during the birds’ moulting season.
An ‘‘island’’, actually the little hill at the tip of a low peninsula connected to
the land east of the Pongola river, is found in the middle of an extensive area of
the Pongola floodplain. Known as Mavilo, its vegetation is, to some extent,
unique. The hill runs N/S with stunted Sand Forest on the east slope while the rest
is rocky and resembles Acacia nigrescens Thicket on boulder beds, with ‘‘pan-
edge’’ species around the base.
Fever Tree Forest
Pure stands of Fever trees, Acacia xanthophloea, occur at the south western
end of Shokwe (a fairly newly formed forest) and at the eastern end of Banzi pan.
This is tall, pure A. xanthophloea, with a uniform crown height of 20 m, with
occasional thickets of Gardenia cornuta, Azima tetracantha, Acacia schweinfurthii
and Maytenus heterophylla. Chloris gayana is the dominant and almost the only
understorey species. The fever trees around Inyamiti form a narrow band and are
thus considered part of the ‘‘pan-edge’’ community.
““Pan-edge’’ Communities
These communities are very variable. Acacia robusta, A.
Sycamorus (missing from Inyamiti, perhaps because of the brackish water)
xanthophloea, Ficus
6 Journal of South African Botany
Sideroxylon inerme, Spirostachys africana and Schotia brachypetala are amongst
the species found growing as thicket, forest or scattered trees. Phragmites spp. and
Cyperus spp. can form an intermediate strip between water, mudflats and trees
(sometimes very extensive.) A separate and quite different area of floodplain or
‘“‘pan-edge”’ is found at both ends of Inyamiti. These are the flats that are exposed
all winter (under normal flood conditions) and flooded for some time each
summer. When the water drops it exposes lawns of Cynodon dactylon with some
Sporobolus virginicus and Digitaria swazica. This area is well utilised by hippo,
rhino, antelope and geese. Inyamiti is brack and produces a salty crust at the
western end. Salicornia pachystachya and Arthrocnemum natalense var. natalense
are an unusual feature of the area.
There have been consecutive, abnormally high rainy seasons which have
caused changes to the margins of some of the pans. Inyamiti and iHotwe pans on
the Pongola floodplain have been affected by high flood levels. The Phragmites
spp. and Nymphaea spp. of Skehla pan (on north bank of Inyamiti) have almost
disappeared, and Acacia xanthophloea is dying off in large numbers, and only
very few young plants occur along the shoreline. Shokwe has been inundated
much longer than usual (starting in December, 1971 and particularly since 1974)
and the A. xanthophloea and the fairly newly established Riverine Forest on the
west bank are suffering considerable numbers of die-offs.
There used to be a small stream flowing from Banzi pan, through the A.
xanthophloea forest into a swamp and eventually draining into the Pongola river.
High floodwaters from Banzi have washed beds of Typha latifolia, Phragmites
spp. and the gently sloping grassy banks away creating a donga and transforming
the area.
Inyamiti pan, famous for the wealth of bird and animal life it supports, is fed
by a stream which drains off the Ndumu farm lands. It is not known how the
insecticides which also drain off, have and will affect the plant and animal life in
and around the pan.
Aquatic communities
The semi-permanent pans of the game reserve and the Pongola river, espe-
cially the old course, hold a wealth of submerged and emergent aquatic plant life.
Floating rafts of a mixture of Ricciocarpos natans, Azolla pinnata, Spirodela spp.
Wolffia arrhiza and Pistia stratiotes are found on the Pongola river and floodplain.
The pans hold changing communities of Potamogeton spp., Ruppia maritima,
Ceratophyllum demersum, Utricularia inflexa var. inflexa and others, which
provide a plentiful supply of food for waterfowl and fish.
In Banzi pan, the once nearly continuous floating raft of vegetation comprising
a mixture of Leersia hexandra, Pycreus mundtii, Oxycaryum cubense (= Scirpus
cubensis), Echinochloa pyramidalis (prostrate form), Trapa natans, Nymphaea
spp., Pistia stratiotes and Ludwigia octovalvis subsp. octovalvis has apparently
See ——— ee — ee a
Checklist of Plants in Ndumu Game Reserve 7
been completely disrupted (1976) and, this being the case, one wonders what has
become of the submerged aquatics. High floodwaters are considered the cause of
the disruption.
The small seasonal pools and water courses of the Mahemane and the west
have some most interesting aquatics, many only found here, such as Oftelia
exserta, Lagarosiphon verticilliflorum, Aponogeton junceus, Burnata enneandra,
Lymnophytum obtusifolium, Nymphoides sp. c.f. rautenanii, Utricularia inflexa
var. stellaris, Scirpus articulatus and Eleocharis fistulosa.
Sand Forest
There are two areas of Sand Forest on Ndumu Hill; Ulukondo and Engabateni,
both of which have suffered at times from over-utilisation by game, particularly
nyala and bushpigs, and disturbance by man. Engabateni has been disrupted to a
greater extent than Ulukondo. East of the Pongola Floodplain there are other
patches of Sand Forest, some fairly extensive, which have a much higher canopy
but are otherwise composed of the same species. The open areas between these
forest patches have a mixture of Ndumu Hill broadleaf species and Sand Forest
species, with grass and occasional thorny thickets on previously disturbed ground.
Newtonia hildebrandtii grows to 12 m and more, emergent above the 6 to 9m
canopy of the forest which has Cleistanthus schlechteri and Pteleopsis myrt:folia
as the dominants, with Brachylaena huillensis, Albizia petersiana subsp. evansit,
A. forbesii, Croton gratissimus, Wrightia natalensis, Hymenocardia ulmoides,
Combretum spp., Strychnos decussata, Haplocoelum gallense and Manilkara
discolor as the taller tree species. Small trees include Monodora junodii, Canthium
spp., Boscia foetida subsp. longipedicellata, Toddaliopsis bremekampu, Gardenia
amoena, Drypetes arguta, Cola microcarpa, Erythroxylon emarginatum, Craibea
zimmermannii, Suregada zanzibarica, Salacia leptoclada and Sapium integerri-
mum. The shrub layer includes Tricalysia spp., Carissa tetramera, Cussonia
arenicola, Dalbergia sp. c.f. D. nitidula, Croton steenkampiana, C. pseudopul-
chellus and Schlerochiton coeruleus. Climbers include Quisqualis parviflora,
Dioscorea cotinifolia, Strophanthus spp., Asparagus spp., Secamone spp., and
Ipomoea cairica. The ground layer is sparse with Stylochiton natalense, Col-
eotrype natalensis, Sansevieria spp., and Panicum deustum.
Acacia tortilis Woodland
There are a number of pure stands of Acacia tortilis. Some of those on the
Mahemane flats area have a canopy of 6 m. Those on the edge of the Usutu
Floodplain have a canopy of 9 m and have many climbers and some thickets, with
good grass cover as an understorey. These plants include the climbers Cocculus
hirsutus, Capparis tomentosa, Diospyros villosa, Sarcostemma viminale, Tylo-
phora lycioides and Senecio pleistocephalus, small trees and shrubs such as
Gardenia cornuta and Azima tetracantha, and a few tall trees such as Trichilia
8 Journal of South African Botany
emetica, Balanites maughamii and Kigelia africana, with the grass Panicum
maximum the dominant herbaceous understorey.
There are large disturbed areas in the west where A. tortilis predominates in
the tree stratum, with thorny scrub and some thicket as understorey, and a sparse
grass fieldlayer. Albizia petersiana subsp. evansii, Acacia grandicornuta, Gar-
denia cornuta, Dichrostachys cinerea, Ehretia rigida, Croton menyhartii, Azima
tetracantha, Aloe marlothii, Capparis citriifolia and Asparagus spp., are among
the more common scrub and thicket-forming plants.
Mahemane Thicket
This covers a great area of the flats and is a dense thicket with Acacia
grandicornuta, Albizia petersiana subsp. evansii and A. anthelmentica predom-
inating in the tree stratum. The area west of Inyamiti has a canopy averaging 6
to 9 m, elsewhere the height of this type of thicket is variable, as is the species
composition. The common larger trees are Balanites maughamii, Pappea capensis,
Manilkara mochisia, Spirostachys africana and Ptaeroxylon obliquum, with scat-
tered Newtonia hildebrandtii. Smaller common trees are Gardenia cornuta,
Maytenus heterophylla, Ehretia rigida, Dichrostachys cinerea, Acacia luederitzii
var. retinens, Cordia ovalis and Euclea spp., with Croton menyhartii, Euphorbia
grandicornis, Carissa spp., Asparagus spp., Ximenia caffra and Senecio barber-
tonicus in the shrub layer. The grasses are sparse in dry seasons, but with good
rains Panicum maximum, P. deustum, Enteropogon monostachyos subsp. afri-
canus and Sporobolus smutsii are common.
Throughout the west and on the lower slopes of Ndumu Hill, there are varying
formations of the plant species found in the Mahemane.
A feature of the area is the small seasonal pools described under Aquatic
Communities.
Deciduous broad-leaf woodland
The Hutton red sands of Ndumu Hill carry deciduous broad-leaf open wood-
land with small scattered thickets, some with Sand Forest species. Sclerocarya
caffra up to 12 m tall, with Acacia burkei and Terminalia sericea predominate
(canopy at 6 to 10 m). Strychnos spinosa, Strychnos madagascariensis, Combre-
tum spp. (5), Albizia petersiana subsp. evansii, Ziziphus mucronata, Dialium
schlechteri, Lannea stuhlmannii, Peltophorum africanum, Sterculia rogersii, Ozo-
roa obovata, Vangueria cyanescens, Rhus gueinzii and Xeromphis obovata are all
frequent tree species. Eugenia mossambicensis, Salacia kraussti and Dichapetalum
cymosum are very common small shrubs forming large communities. Cassia
petersiana, usually growing as a shrub is on the increase and has spread rapidly in
small thickets. The grass cover is generally good, up to 1 m high, and attracts
large numbers of animals to the hill. Panicum maximum, Pogonarthria squarrosa,
Aristida spp., Digitaria pentzii, Eragrostis superba, E. rigidior, Sporobolus
Checklist of Plants in Ndumu Game Reserve 9
fimbriatus, Perotis patens, Eustachys paspaloides, Heteropogon contortus,
Schmidtia pappophoroides and Tricholaene monachne are common while Hyper-
thelia dissoluta is becoming more widespread.
On the NW slopes, after a number of consecutive seasons with good rains, the
drainage lines have formed a “‘sponge’’ (which is dissected by the main road)
from which tiny streams strickle down towards Inyamiti. A different community
of plants, favouring the very wet conditions, can be found here.
Acacia nigrescens Woodland
On the eastern slopes of Ndumu Hill and roughly encircling it on old boulder
beds there is Acacia nigrescens Thicket. The other larger trees are Combretum
spp., Schotia brachypetala, Albizia spp., Acacia gerrardii, A. burkei, Galpinia
transvaalica, Bolusanthus speciosus, Peltophorum africanum and Berchemia zey-
heri. Smaller trees such as Ehretia rigida, Rhus gueinzii, Dichrostachys cinerea,
and shrubs such as Grewia caffra, G. villosa, and Croton menyhartii also occur.
The drainage lines have slightly different species, Spirostachys africana and
Sideroxylon inerme among them. Panicum maximum, Heteropogon contortus and
Themeda triandra are amongst the most common grasses.
In the west, there is a mosaic of woodland and savanna areas interspersed with
shrub and thicket. There are almost pure stands of Acacia nigrescens Woodland
with scattered large specimens of Combretum imberbe, Lonchocarpus capassa and
Bolusanthus speciosus. Themeda triandra predominates in the 1 m high grass
cover. A feature of the area is the circular grass formations around termitaria. The
main constituents of these communities are Aristida congesta subsp. barbicollis,
Heteropogon contortus and Digitaria argyrograpta, but these grasses have not
been adequately collected and it is possible that more species might be present.
Evenly spaced Acacia nilotica Woodland with occasional A. nigrescens covers
a large area. Panicum maximum and Themeda triandra provide the good grass
cover.
A. nilotica and A. luederitzii var. retinens on wet soils near drainage lines in
the west, form a fairly uniform low woodland with A. senegal, Dichrostachys
cinerea, Albizia petersiana subsp. evansii forming thickets. Grasses are Panicum
maximum, P. coloratum and Urochloa mossambicensis.
In an area west of the Mahemane, where Chloris roxburghiana is fairly
common, A. nigrescens is the dominant tree species, and with Ziziphus mucronata,
Albizia petersiana subsp. evansii, A. grandicornuta, forms an open woodland with
scattered thickets of Cordia ovalis and Croton menyhartii.
There are well-defined seasonal water courses in the west. One of these drains
into the western end of Inyamiti pan after passing through the Balemhlanga
swamp on the south boundary fence-line. The other flows through from the
south-western corner of the reserve, eventually draining into the Usutu floodplain.
10 Journal of South African Botany
Some species commonly found on the Lebombo can be found here, namely
Heteropyxis natalensis, Bauhinia galpinii and Vitex harveyana.
The rhyolite outcrops in the Mkonjane area are unique in the reserve and,
although a very small part of it, are an important protected example of the
Lebombo foothills. Pavetta edentula, Acacia caffra, Olax dissitiflora, Pterocarpus
rotundifolius and Combretum apiculatum are typical Lebombo mountain species.
Mountain and rhyolite grasses and sedge species found here are Diheteropogon
amplectens, Sporobolus festivus var. fibrosus, Tripogon minimus, Oropetium
capense and Mariscus sp. prob. M. dubius.
TABLE 2.
A LIST OF ANGIOSPERM FAMILIES COMPRISING ONE PER CENT AND MORE OF
THE TOTAL NUMBER OF SPECIES.
NUMBER OF NUMBER OF
FAMILY GENERA SPECIES as % of TOTAL
POACEAE (Graminae) ....... Sil 13,0
PEGUMINO SAB eee 35 9,1
COMEOSIRAES See 29 6,3
EUPHORBIACEAE SAE ae: 20 4,9
PIBINCE AB eee 15 3,9
RUBIACEAE ear 19 3,9
GYEPBRAGCE AR EE 10 3,8
NIANIWIANGE AR eee 8 DA
ACANTHACEAE aeons 13 2,5
ASCEERIADAGEAE Sse eae 14 De)
CONVORVULAGCEAE = 7 DB
LAMIACEAE (Labiatae)...... 12 2,0
AMARANTHACEAE ........ 13 7
CAPPARACEAE oe 7 Lod
CUGCURBIMACEAES eee 10 17
VERBENACEAE EEE 10 1.5
COMBREDACBAE == 55.552 4 1,5
COMMELINACEAE ......... 6 133
INRO@GYNAGE ARS 9 1,3
TILIACEAE Rare ee, Shere 3 le,
LORANTHACEAE .......... 2 il. Jl
BORAGINACEAE .......... 4 itl
SOLANACEAE Reece 4 1,0
CELPASTRAGEAES Seer 5 1,0
ORCHIDACEAE eer 6 1,0
FLORISTICS
Analysis of the flora of Ndumu Game Reserve, based on this checklist, reveals
the following plant species: 1 Bryophyte, 4 Pteridophytes, 241 Monocotyledons
and 644 Dicotyledons. Angiosperm families comprising | per cent and more of the
total number of species are listed in order of numerical importance in Table 2.
Varieties, sub-species and forms have not been counted unless the record is the
Checklist of Plants in Ndumu Game Reserve 11
only one for the species to which they refer (if these were to be included the
checklist would number over 900 entities).
The largest genera are:
Cyperus, Acacia (15); Eragrostis (13); Hibiscus, Ipomoea (11); Digitaria,
Asparagus, Euphorbia, Senecio (10); Combretum (9); Loranthus, Indigofera,
Helichrysum (8); Panicum, Sporobolus, Grewia, Pavetia, (7); Phyllanthus, Plec-
tranthus, Justicia, Vernonia (6); Setaria, Commelina, Aloe, Albizia, Cassia,
Crotalaria, Ceropegia, Heliotropium, Solanum (5).
CHECKLIST
The arrangement follows that of the Flora of Natal (Ross, 1972), and where
additional material is represented it follows the Genera of Southern African
Flowering Plants (Dyer, 1976). The ferns are arranged according to Flora
Zambesiaca, Pteridophyta (Schelpe, 1970). The numbers in brackets before family
names are those used in Ross, and the generic number, from the same source, is
listed in the left hand margin next to the first species of the genus.
All herbarium numbers following the specific names are mine except where
another collector’s initial is attached to the number (an explanation of the initials
is listed below). Where no number is supplied, the plant is well known and has
been observed but not collected. An asterisk (*) indicates that a plant occurs in the
camp or crocodile farm gardens and was transplanted from outside, but nearby the
game reserve. Only one collector’s number has been listed for each plant although
there are often many more specimens in the herbarium.
Some notes on the plants have been included but are entirely random
observations, made in Ndumu Game Reserve. Many of the plants must be utilised
to a far greater extent by insects, animals and man, but only known personal
observations (with some additions from A. C. Pooley) and where indicated, from
the herbarium notes of other collectors in the reserve, have been listed. Some
information was also extracted from various papers such as Anderson and Pooley
(1977), Dixon (1966), Dutton (1972), Oatley (1964), Pooley (1968), Scotcher
(1974) and Tinley (1964a).
Some Zulu names of plants with edible fruits have been included, but no
attempt has been made to provide a complete list of Zulu names.
Such observations on locality and abundance of plants as are included, are
subject to the effect of rainfall, the artificial and irregular flooding of the Pongola
river caused by the Jozini dam, and the feeding and trampling effect of the animal
populations within the reserve. Reference to locality of plants indicates the area in
‘which they are most commonly found, and are not the only area in which they are
found.
12 Journal of South African Botany
Growth form (after Compton, 1966) has been described in the broadest terms,
to assist the field worker, and these are defined as:
Trees: a typical tree is a woody plant with a single main trunk. However, some
multistemmed woody plants are too well developed to be called shrubs and are
listed as trees.
Shrubs: mostly woody plants which do not usually produce a single main trunk,
branching from ground level, with many more-or-less equivalent and often
interlacing branches. Some single stemmed plants are of small stature and limited
development, and have, therefore, been classed as shrubs.
Herbs: this includes a large number of growth forms in which stems are relatively
soft, although the lower parts of stems and rootstock can be woody.
Bulbs: perennial plants whose underground resting organs have a swollen form
and act as storage receptacles (bulbs, corms, rhizomes).
Succulents: this group includes a variety of plants whose most conspicuous feature
is a fleshy thickening of the aerial parts.
Parasites: plants, partially or wholly dependent on the host plant for nutrition.
Epiphytes: plants independent of soil in any quantity and which can absorb water
and nourishment from plant debris through clasping and absorbent roots.
Aquatics: includes plants dependent on open water, either free floating, rooted in
underwater-mud with flowers and leaves on surface, or completely submerged.
PPdM P. P. de Moor
HF H. Furness
JVH J. V. Hancock
H&B Hilliard and Burtt
EJM E. J. Moll
ICP I. C. Player
JHR J. H. Ross
JSBS J. S. B. Scotcher
KLT K. L. Tinley
T&W Tinley and Ward
CJW C. J. Ward
BRYOPHYTA
HEPATICAE
RICCIACEAE
Ricciocarpus natans (L.) Corda 1653 a Aquatic. Pongola Floodplain; flooded
depressions.
Checklist of Plants in Ndumu Game Reserve 13
PTERIDOPHYTA
EQUISETALES
Equisetum ramosissimum Desf. 1355 Herb. Usutu Floodplain. Frequent on
sandy alluvium, amongst grasses.
FILICALES
MARSILIACEAE
Marsilia apposita Launert 1599 Aquatic in seasonal pools. Mahemane.
AZOLLACEAE
Azolla pinnata R.Br. 1653 Aquatic. Plentiful in Pongola river.
ADIANTACEAE
Pellaea viridis (Forsk.) Prantl 486 Herb. Ulukondo. Sand Forest.
THELYPTERIDACEAE
Thelypteris dentata (Forsk.) E. St. John 487 Herb. Usutu Riverine Forest, in
damp depressions.
SPERMATOPHYTA
MONOCOTYLEDONEAE
(5) TYPHACEAE
49 Typha latifolia L. subsp. capensis Rohrb. 1484 Isolated patches on flood-
plains, not widespread. Favoured by Thick-billed weaver birds for their nests,
which are usually constructed between two stems.
(7) POTAMOGETONACEAE
58 Potamogeton crispus L. 629 Aquatic. Inyamiti, Banzi Pan, common. Valuable
source of food for waterfowl and fish and provides shelter and food for molluscs
and aquatic insects.
P. pectinatus L. 491 KLT Aquatic. Inyamiti pan. Valuable source of food for
waterfowl and fish and provides shelter and food for molluscs and aquatic insects.
(8) RUPPIACEAE
59 Ruppia maritima L. 507 KLT Aquatic. Inyamiti pan.
(10) NAJADACEAE ck
64 Najas marina L. subsp. delilei (Rouy) Maire 3208 CJW Aquatic. Inyamiti
pan. Source of food for wildfowl and fish. ;
N. pectinata (Parl.) Magnus 457 KLT Aquatic. Banzi pan.
(11) APONOGETONACEAE ;
65 Aponogeton junceus Lehm. ex Schlechtd. 1577 Aquatic. Uncommon in sea-
sonal pools in Mahemane.
(13) ALISMATACEAE '
72 Lymnophytum obtusifolium (L.) Mig. 1575 Aquatic herb. Frequent in seasonal
pools in Mahemane. : : | ine
Burnata enneandra Micheli 1579 Herb. Fairly frequent in seasonal pools In
Mahemane (1st record for Natal).
14
(14)
88
95
(15)
107
119
125
126
127a
132
Journal of South African Botany
HYDROCHARITACEAE
Lagarosiphon verticillifolius Oberm. 1551 Aquatic. Frequent in seasonal pools
in Mahemane.
Ottelia exserta (Ridley) Dandy 1548 Aquatic. Uncommon in seasonal pools in
Mahemane. Growing in | m of water, leaves floating, flowers erect.
PGACEAE (Nom. alt. Gramineae)
Coix lacryma-jobi L. 589 Uncommon. Usutu Floodplain. The nut-like, hard
sheaths of inflorescences are used by Tongas as beads.
Ischaemum brachyatherum (Hochst.) Fenzl 1095 Edge of Pongola Floodplain.
Urelytrum squarrosum Hack. 1216 Ndumu Hill. Uncommon in deciduous
broad-leaf woodland.
Rhytachne robusta Stapf 895 KLT On drainage lines and in the Balemhlanga
swamp.
Hemarthria altissima (Poir.) Stapf & C. E. Hubb. 1163 Floodplains. Exten-
sively grazed by hippo and antelope.
Elyonurus argenteus Nees KLY Common in A. nigrescens Woodland.
134 Andropogon gayanus Kunth var. squamulatus (Hochst. ex A. Rich.),
134A
134D
134G
134J
134N
134R
136
Stapf 1232 Ndumu Hill. Frequent in deciduous broad-leaf woodland.
Sorghum halepense (L.) Pers. 402 KLT
S. versicolor Anderss. 1618 Mkonjane. Uncommon in A. nigrescens Wood-
land.
S. verticilliflorum (Steud.) Stapf 1427 Common on floodplains. New leaves
after burn grazed by hippo. Vervet monkeys eat seeds.
Bothriochloa glabra (Roxb.) A. Camus 1569 Usutu Floodplain. Seasonally
flooded depressions.
B. insculpta (Hochst.) A.Camus 781KLT ‘‘Invaded large areas of Themeda veld
on black loam soils’’ (KLT).
Cymbopogon excavatus (Hochst.) Stapf ex Burtt Davy 1188 Usutu Floodplain.
Common.
C. plurinodis (Stapf) Stapf ex Burtt Davy 868 KLT ‘Acacia nigrescens veld”
(KLT).
C. validus Stapf ex Burtt Davy 1318 Scattered communities on Usutu Flood-
plain.
Heteropogon contortus (L.) Beauv. ex Roem. & Schult. 1047 Widespread on
floodplains, in woodland and a constituent of the circular grass communities
around termitaria in the west.
Diheteropogon amplectens (Nees) W. D. Clayton 1023 KLT Mkonjane. Com-
mon on sandy soil of rhyolite outcrops.
Hyperthelia dissoluta (Nees ex Steud.) W. D. Clayton 558 Usutu Floodplain
and common on Ndumu Hill in deciduous broad-leaf woodland. Sometimes cut
for thatch for the game reserve. The area under H. dissoluta on Ndumu Hill has
increased considerably and there is cause for speculation as to whether cutting has
aggravated the position.
Themeda triandra Forsk. var. trachyspathea Goossens 1319 Widespread;
dominating in Acacia nigrescens Woodland. Grazed by hippo, rhino and antelope.
143
148
161
164
166
166B
166C
166D
166E
166F
Checklist of Plants in Ndumu Game Reserve 15
Tragus berteronianus Schult. 1223 Widespread. Common on disturbed areas.
Perotis patens Gandoger 562 Ndumu Hill. Frequent on Hutton red sands in
deciduous broad-leaf woodland.
Paspalum commersonii Lam. 1179 Floodplains, above flood water level.
Hippo and antelope graze it.
P. urvillei Steud. 93 JSBS
P. vaginatum Swartz 1162 Inyamiti pan (an escape from cultivation). Com-
mon on edges of pans, forming pure stands.
Eriochloa meyeranum (Nees) Pilg. (syn. Panicum meyerianum Nees, also included
is P. meyeranum Nees var. grandiglume Stent & Rattray KLT
508) 578 Grazed by hippo, rhino and antelope. Common on floodplains.
E. nubica (Steud.) Stapf 1244 Inyamiti pan margins. Common on Usutu
Floodplain above Phragmites line.
E. stapfiana Clayton 1370 Balemhlanga. Common in seasonal pools.
Panicum chusqueoides Hack. 1397 E of Pongola Floodplain, in shade of Sand
Forest.
P. coloratum L. 1243 Floodplains, seasonal pools. Grazed by hippo, antelope.
P. deustum Thunb. 1249 Common in Mahemane and Ulukondo—in dense
thicket and in Sand Forest. Grazed.
P. infestum Anderss. ex Peters 1592 Ndumu Hill drainage line.
P. kalaharense Mez 1217 Ndumu Hill. Infrequently scattered in deciduous
broad-leaf woodland.
P. maximum Jacg. 559 Widespread and common especially in woodland on
Ndumu Hill and in pure stands of A. tortilis. Important grazing for hippo, rhino
and antelope.
P. subalbidum Kunth. (syn. P. glabrescens Steud.) 2945 CJW Floodplain.
‘“*Common in wet mud’’ (KLT).
Urochloa mosambicensis (Hack.) Dandy 1254 Widespread; common on dis-
turbed areas. Grazed by hippo and antelope.
U. trichopus (Hochst.) Stapf 962 KLT Usutu Floodplain.
Brachiaria nigropedata (Munro) Stapf 1563 b Ndumu Hill drainage line.
B. xantholeuca (Hack.) Stapf. 1563a N slope, Ndumu Hill. Woodland.
Echinochloa colona (L.) Link. 1457 Pongola Floodplain, on edge of water.
E. holubii (Stapf) Stapf 1262 Common in seasonally inundated pools and
depressions. ave my
E. pyramidalis (Lam.) Hitchcock & Chase This species has two distinct growt
forms:
1354 *‘Funga makala’’—‘‘erect’’ form occurs on soil that dries out after summer
floods. This form is utilized very little by animals (“‘hairy form” Ward 2030
according to Flora of Natal). : ; :
1353 *‘Mbuku’’—‘‘prostrate’’ form occurs in wet depressions on the floodplain
or actually floats at the edges of permanent or semi-permanent _ pans. It is
extensively grazed, particularly by hippo (“glabrous form’’ Ward 6151 according
to Flora of Natal).
Sacciolepis curvata (L.) Chase 1221
woodland and Sand Forest.
Digitaria adscendens (H.B.K.) Henr. 1384 Ndumu Hill. Deciduous broad-leaf
woodland.
D. argyrograpta (Nees) Stapf
munities around termitaria in the we
Ndumu Hill. In deciduous broad-leaf
1379 A constituent of the circular grass com-
stern A. nigrescens Woodland.
168
168A
169
17
—
174
175
194
Journal of South African Botany
Digitaria eriantha Steud. 1937 JHR Ndumu Hill. Deciduous broad-leaf wood-
land.
D. gymnostachys Pilg. 1651 Ulukondo. Growing on white sand in Sand
Forest.
D. longiflora (Retz.) Pers. 1587 Ndumu Hill, on seasonally inundated
‘sponge’ near main gate.
D. pentzii Stent 1371 Widespread in woodland and on dry floodplain. Grazed
by hippo and antelope.
D. sp. c.f. smutsti Stent 54 ICP Inyamiti pan margins. Grazed by hippo.
Almost impossible to distinguish from D. macroglossa Henr.
D. swazilandensis Stent 1627 Floodplains and seasonal pools. Grazed by
hippo.
D. ternata (A. Rich.) Stapf 56 ICP Inyamiti pan margins. Grazed by hippo.
Annual weed.
D. zeyheri (Nees) Henr. 1038 Pongola Floodplain margins. Grazed by hippo.
Tricholaena monachne (Trin.) Stapf & C. E. Hubb. 1226 Usutu Floodplain
and Ndumu Hill, woodland.
Rhynchelytrum repens (Willd.) C. E. Hubb. 1428 Widespread. Common on
margins of roads.
Oplismenus hirtellus (L.) Beauv. JSBS_ Riverine Forest.
Setaria chevalieri Stapf ex Stapf & C. E. Hubb. 395 Riverine Forest. Grazed
by hippo.
S. neglecta de Wit 862 KLT Common in Acacia/Albizia Thicket in western
area.
S. sphacelata (Schumach.) Stapf & C. E. Hubb. ex M. B. Moss 18 PPdM
S. verticillata (L.) Beauv. 1503 Pongola Floodplain.
S. woodii Hack. 1190 Floodplains and seasonally inundated depressions.
Cenchrus ciliaris L. 1805 Disturbed areas.
Pennisetum glaucocladum Stapf & C. E. Hubb. 1426 Infrequent stands on
floodplains near water.
Leersia hexandra Sw. 1580 Banzi pan in floating raft of vegetation over 3 m
of water.
208 Aristida adscensionis L. (syn. A. curvata (Nees) Trin. & Rupr.) 41 PPdM
230
A. canescens Henr. subsp. canescens 617 Usutu Floodplain.
A. congesta R. & S. subsp. congesta 1016 Widespread in woodland and open
grassland. Hippo and antelope graze it.
A. congesta R. & S. subsp. barbicollis (Trin. & Rupr.) de Winter 998 Wide-
spread in woodland and open grassland. Constituent of the circular grass communi-
ties around termitaria in the western Acacia nigrescens veld. Grazed by hippo and
antelope.
A. ne Hack. ex Schinz subsp. graciliflora (Pilger) Melderis 1235 Wide-
spread in woodland. Grazed by hippo and antelope.
Sporobolus consimilis Fres. 1039 KLT Floodplain margins.
S. festivus Hochst. ex A. Rich. var. fibrosus Stapf ex Stent 1017 KLT Uncom-
mon on sandy soil in depressions on rhyolite outcrops. Mkonjane.
S. fimbriatus Nees forma 1081 Usutu Floodplain. Included is S. fimbriatus
Nees var. /atifolius Stent 1300 Common, widespread especially on disturbed
areas. Grazed by hippo and antelope.
S. nitens Stent 1161 Mahemane, near paths and roads. Grazed by hippo and
antelope.
282
287
288
288A
298
299
302
305
307
310
3172
S17.
333
3377)
337A
Checklist of Plants in Ndumu Game Reserve 17
Sporobolus pyramidalis Beauv. 1330 Usutu Floodplain.
S. smutsii Stent 1258 Mahemane. Common in thicket. Grazed by hippo and
antelope.
S. virginicus (L.) Kunth 1160 Inyamiti pan margins. Extensively grazed by
hippo and antelope.
Cynodon dactylon (L.) Pers. 1036 Widespread especially at old kraal sites and
disturbed areas; also around Inyamiti where it forms ‘lawns’ which provide food
for geese, hippo and antelope and probably fish when the grass is inundated.
Enteropogon macrostachyos (Hochst. ex A. Rich.) Munro ex Benth. s.n. JSBS
E. monostachyos (Vahl) K. Schum. ex Engl. subsp. africanus W. D. Clayton
1_ 1 Mahemane Thicket.
Chloris gayana Kunth 587 Floodplains and pan margins. Common. Grazed by
hippo and antelope.
C. roxburghiana Schult. 1306 Western area. Locally common in Acacia
nigrescens /Albizia evansii Woodland, west of Mahemane on S boundary.
C. virgata Sw. 619 Floodplains; Bunguzane; Red Cliffs. Grazed by hippo and
antelope.
Eustachys paspaloides (Vahl) Lanza & Mattei 999 Widespread, but nowhere
abundant.
Tripogon minimus (A. Rich.) Hochst. ex Steud. 1024 KLT Mkonjane, in
shallow depressions on rhyolite outcrops.
Tetrapogon mosambicensis (K. Schum.) L. Chippindall ex B. S. Fisher
1389 Seasonal pools, quite common in west.
Dinebra retroflexa (Vahl) Panzer 474 Pongola Floodplain margins.
Dactyloctenium australe Steud. 1642 Ndumu Hill. Woodland. '
D. geminatum Hack. 34 PPdM May only be a robust form of the closely allied
stoloniferous D. australe.
D. giganteum Fisher et Schweickerdt 1206 Ndumu Hill. Frequent in wood-
land. Grazed. Annual not stoloniferous.
Leptochloa panicea (Retz.) Ohwi 1620 Floodplains. me
L. uniflora A. Rich. (syn. Craspedorachis uniflora (A. Rich.) Chippindall) 1000
KLT Engabateni. In fairly deep shade of Sand Forest.
Enneapogon cenchroides (Licht.) C. E. Hubb. 1084 Lower slopes of Ndumu
Hill in thicket. :
E. scoparius Stapf 1423 E of Pongola Floodplain; Mavilo.
Schmidtia pappophoroides Steud. 552 Ndumu Hill. Common in shade in
woodland.
Triraphis schlechteri Pilger ex Stent 116 JSBS
Phragmites australis (Cay.) Trin. ex Steud. 516 iHotwe and Sabatane. On
floodplains and pan edges. L
P. paiitianus rcniiss 448 Usutu river banks. On floodplains. Monkeys eat
new shoots. Tonga herd boys make flutes from the dried stem of the reed.
Diplachne eleusine Nees 1040 Balemhlanga. Common on drainage lines in
west. Widespread in woodland.
D. fusca (L.) Beauv. ex Stapf s.n. JSBS
Pogonarthria squarrosa (Licht.) Pilg. 554 Ndumu Hill. Woodland.
18
337C
34
—
398
(16)
459
Journal of South African Botany
Trichoneura grandiglumis (Nees) Ekman 555 Ndumu Hill. Woodland.
T. sp. 1650 Common on Sand Forest floor in Engabateni, Ulukondo. White
sand.
Eragrostis aspera (Jacq.) Nees 1377 Ndumu Hill. Woodland.
E. atherstonei Stapf 1041 Ndumu Hill. Woodland.
E. cilianensis (All.) Lutati 161 Ndumu Hill down to Pongola Floodplain and
Inyamiti.
E. ciliaris (L.) R.Br. 613 Widespread in woodland and open grassland.
E. cylindriflora Hochst. (syn. E. horizontalis Peter) 1374 Balemhlanga. Sea-
sonally inundated depression.
E. gummiflua Nees 1394 Ndumu Hill. Woodland.
E. heteromera Stapf 1231 Usutu Floodplain. Widespread, especially near
seasonally inundated pools, and depressions on floodplain. Grazed by hippo.
. Namoena K. Schum. 1045 KLT Inyamiti pan margins.
pallens Hack. 1227 Ndumu Hill. Woodland.
rigidior Pilg. 1234 Ndumu Hill; Usutu Floodplain. Grazed by hippo.
rotifer Rendle 1603 Seasonal pools in west.
sp. c.f. E. stapfii De Winter 1203 Ndumu Hill. Woodland.
E. superba Peyr. 1017 Widespread in woodland and in open grassland on
Usutu Floodplain. Grazed by hippo and antelope.
E.sp. 1327 Usutu Floodplain.
By By By Gr
Oropetium capense Stapf 1027 KLT Common on rhyolite outcrops, in shallow
depressions. Mkonjane.
CYPERACEAE
Note: In this family species are being moved from one genus to another with
what sometimes seems incomplete supporting evidence. Scirpus and to a lesser
extent Bulbostylis are affected. Until the situation is more stabilised, established
generic limits have mostly been followed in this list (Dr. K. D. Gordon-Gray).
Cyperus articulatus L. 1630 Near water’s edge. Banzi pan margins.
C. denudatus L.f. 1566 Seasonal pools.
C. denudatus var. sphaerocarpus (Schrad.) Kuk. 1680 Ndumu Hill drainage
lines.
C. difformis L. 1625 Balemhlanga. Seasonally inundated depressions.
C. digitatus Toxb. subsp. auricomis (Sieber) Kuk. 365a Banzi pan mar-
ins.
C. distans L.f. 365 Banzi pan margins and seasonally inundated depressions.
C. fastigiatus Rottb. 1180 Common on floodplains. Hippo graze. Used for
making sleeping mats.
C. immensus C.B.Cl. 957 KLT Local patches in Mdipine and Ukehla areas, in
reedbeds.
C. margaritaceus Vahl 1012 Ndumu Hill. Woodland.
C. nudicaulis Poir. 1639 In floating ‘raft’ of vegetation on Banzi pan, over 3
m of water.
C. obtusiflorus Vahl 4313 EJM_ Seasonal sponge on main road near main gate.
C. papyrus L. 624 Scattered patches on floodplains, near pans.
C. rotundus L. 1502 Edge of Pongola Floodplain.
C. rupestris Kunth 1586 Drainage line, Ndumu Hill.
C. sexangularis Nees 1430 Mdipine area. Seasonally flooded depression on
Usutu Floodplain.
C. teneriffae Poir. 1166 Scattered in Mahemane Thicket.
Checklist of Plants in Ndumu Game Reserve 19
Oxycaryum cubense (Poep. & Kunth) K. Lye (syn. Scirpus cubensis Poeppig &
en ex Kunth) 1471 Banzi pan. In floating raft of vegetation, over 3 m depth
of water.
459A Pycreus mundtii Nees 1470 Banzi pan. In floating raft of vegetation, over 3 m
of water.
P. pelophilus (Ridl.) C.B.Cl. 1643 On sponge on main road, Ndumu Hill.
P. polystachyos Beauv. 1046 KLT Inyamiti pan.
459C Mariscus sp. c.f. M. capensis Schrad. 1591 Ndumu Hill drainage line.
M. sp. prob. dubius (Rottb.) Hutch. 1468 In depressions on rhyolite outcrops.
Mkonjane.
M. sp. c.f. indecorus (Kunth) Podlech 1593 a & b Ndumu Hill drainage line.
462 Kyllinga alba Nees 685 JHR Ndumu Hill and Acacia nigrescens Woodland on
slopes of Ndumu Hill.
K. erecta Schum. 687 JHR Ndumu Hill. Woodland.
467 Fuirena pubescens (Poir.) Kunth 16 PPdM
F. leptostachya Oliv. 1644 Annual. On sponge on main road, Ndumu Hill.
468 Scirpus articulatus L. 1623 Balemhlanga. Locally common, in seasonal pools.
S. littoralis Schrad. 1682 Inyamiti pan, in 2 m of water. Used for making
sitting mats.
S. muricinux C.B.Cl. 1263 Mahemane. Frequent in seasonal pools.
469 Eleocharis fistulosa Link 1624 Balemhlanga. Frequent in seasonal pools.
471 Fimbristylis bis-umbellata (Forsk.) Bub. 840 KLT Usutu Floodplain. Locally
abundant.
—
471A Bulbostylis burchellii (Fic. & Hiern) C.B.Cl. 611 Usutu Floodplain, Ndumu
Hill. Open grassland and woodland.
B. contexta (Nees) Bodard 1943 JHR Ndumu Hill. Woodland.
B. parvinux C.B.C1. 2015 CJW Ndumu Hill. “Common throughout sandy
soil community’? (CJW)
(17) ARECACEAE (nom. alt. Palmae)
528 Phoenix reclinata Jacq. 59/33 PPdM Common along water courses. Seldom
growing as tall as at coast. Fruits eaten by bushpig, antelope, monkeys and man
(iSundu). Leaves stripped by weaver birds for nests. The stem was tapped in
Ndumu for the sweet sap which is used to brew beer. The young fruits attract
many insects by day (ants, wasps, bees) and many moths by night. It is difficult to
observe what they are all feeding on—they appear to be sucking sap.
553 Hyphaene natalensis Kuntze 609 Ndumu Hill. Uncommon. Woodland. The
fibre from the leaves is used for binding and in the manufacture of baskets (iLala).
(18) ARACEAE .
692 Zamioculcas zamiifolia (Lodd.) Engl. 1638 Bulb. Ulukondo. Common in Sand
Forest.
693 Gonatopus sp. 322 Bulb. Widespread—occasionally in large communities.
Flaweting Beal needed. In woodland, thicket and Sand Forest. Bushpig eat the
whole plant.
764 Stylochiton natalense (Sond.) Schott 741 Bulb. Ndumu Hiill. commen in
woodland and in Sand Forest where it grows as a much more robust plant.
20
791
(19)
794
796
(24)
896
Journal of South African Botany
Pistia stratiotes L. 681 Aquatic. Banzi pan; Mvutsheni pan; Pongola river
“Old Course’. Forms large floating mats.
LEMNACEAE
Spirodela polyrhiza (L.) Schleid. Aquatic. Common constituent of aquatic
floating plant communities on Pongola Floodplain (Musil).
S. punctata (G. F. W. Meyer) Thompson Aquatic. Common constituent of
aquatic floating communities on Pongola Floodplain (Musil).
Wolffia arrhiza (L.) Horkel ex Wimmer Aquatic. Common constituent of
aquatic floating communities on Pongola Floodplain (Musil).
COMMELINACEAE
Commelina africana L. 1004 Herb. Common, widespread. Flowers close at
noon.
C. benghalensis L. 1265 Herb. Common, Ndumu Hill. Woodland. Flowers
close at mid-morning.
C. diffusa Burm.f. 1360 Herb. Common on Pongola Floodplain. Almost
exclusive herb in Cyperus fastigiatus area of Pongola Floodplain on black clay
exposed after flooding. Flowers close at noon.
C. erecta L. 412 Herb. Common, Ndumu Hi!l. Woodland and Riverine Forest.
C. forskalaei Vahl 1634 Herb. Ndumu Hill. Common. Woodland (1st record
for Natal).
899 Aneilema dregeanum Kunth 1399 Common east of Pongola Floodplain in low
899A
899B
903
904
(27)
963
975A
985
scrub. Flowers close at night.
A. petersii (Hassk.) C.B.Cl. 1808 Herb. Frequent in shade. Ndumu Hill.
Woodland.
A. schlechteri K. Schum. 1679 E slope Ndumu Hill. Large communities in
woodland. Grazed.
Murdannia simplex (Vahl) Brenan 202 Herb. Ndumu Hill, on drainage lines.
Flowers open mid-afternoon, close at sunset. Grow in small clumps. Grazed by
antelope.
Ballya zebrina (Chiov.) Brenan 1255 Prostrate herb. Mahemane. Common in
thicket.
Coleotrype natalensis C.B.C1. 287 Herb. Ulukondo, common locally in Sand
Forest. Open early morning, close late afternoon.
Cyanotis speciosa (L.f.) Hassk. 134 Bulb. Ndumu Hill slopes. Flowers close
at midday. When flowers close, stamens curl inwards, pollen first, one over
another, then the petals.
LILIACEAE
Gloriosa superba L. 1825 Bulb, climber with tendrils. Ndumu Hill. Frequent,
especially on the edge of thickets where it is protected from browsing antelope.
Camptorrhiza strumosa (Bak.) Oberm. 915 Bulb. Scattered communities on
Ndumu Hill. Grows in full sun amongst grasses in woodland.
Bulbine asphodeloides (L.) Roem. & Schult. 37 Herb. Mkonjane, margins of
Pongola Floodplain. In Acacia nigrescens Woodland. Flowers close at night, not
to re-open.
989 Anthericum galpinii Bak. var. galpinii 246 Bulb. Ndumu Hill. Frequent,
especially on slopes. Woodland. Flowers open early morning, close midday.
Checklist of Plants in Ndumu Game Reserve 21
989A Trachyandra saltii (Bak.) Oberm. var. secunda (Krause & Dinter) Oberm.
229 Herb. Common on Ndumu Hill. Woodland. Flowers open early afternoon
and close late evening. Grazed.
990 Chlorophytum sp. c.f. C. aridum Oberm. 1420 Herb. Mavilo. In shade of
thicket on ‘island’ in swamps.
C. comosum (Thunb.) Jacques 497 Herb. Widespread in Mahemane Thicket.
Browsed by antelope and rootstock eaten by bushpig.
1012 Eriospermum mackenii (Hook.f.) Bak. 131 Bulb. On seepage line, Ndumu
Hill; Mkonjane. Leaves curl up at sides in heat of day. Flowers open towards
midday and close after dark. Flower after rain, a number of times a year.
1026 Aloe chabaudii Schonl. 634 Succulent. Common in Ulukondo. Sand Forest.
Leaves and flowering stems browsed.
A. marlothii Berger var. marlothii 643 Succulent. Widespread, common, espe-
cially in disturbed areas and in Acacia nigrescens Woodland. Monkeys, birds,
insects (especially bees, ants, flies, beetles) and people suck the nectar. Dry old
leaves are used by the Tongas in making snuff.
A. parvibracteata Schonl. var. zuluensis (Reynolds) Reynolds 644 Succulent.
Widespread, common, sometimes forming large communities. Monkeys eat
flower buds. Leaves and flowers browsed by rhino and antelope.
A. rupestris Bak. Succulent. Ulukondo. Rare. In Sand Forests. Large camp
garden specimens transplanted from Ulukondo in early 1960s.
A. sessiliflora Pole Evans * Succulent. From Abercorn drift track, on
Lebombo foothills just outside game reserve western boundary. Planted in camp
and crocodile farm gardens.
1079 Albuca angolensis Welw. 220 Bulb. Mahemane, Ulukondo. In large communi-
ties in thicket and Sand Forest.
A. pachychlamys Bak. 663 Bulb. Mkonjane. In open woodland.
1080 Urginea delagoensis Bak. 642 Bulb. Mahemane. Frequent but scattered in
thicket. Browsed by antelope. rai
U. epigea R. A. Dyer 3 Bulb. Ndumu Hill. In Acacia nigrescens Woodland.
Scattered communities. Plants form clumps. Flowers open early morning, close
by early afternoon. Browsed by antelope.
U. indica (Roxb.) Kunth var. multiflora Oberm. 661 Bulb. Mahemane. Fre-
quent in thicket. Flowers open at about 13h30, close in evening. Common but
widely scattered. Leaves and flowers browsed.
U. indica (Roxb.) Kunth (query) 127 Bulb. Ndumu Hill. This plant differs
markedly from 661—has different bulb and leaf; fewer, larger flowers; 1s heavily
and unpleasantly scented (661 is unscented); flowers at night (661 opens at
midday, closes in the evening). Numerous specimens of both plants occur in the
same locality on the margins of Pongola Floodplain and lower Ndumu Hill near
the crocodile farm. 127 is common on the sponge on the main road near the main
gate. Buds droop downwards, flowers open upwards.
1084 Dipcadi marlothii Engl. 1583 Bulb. Widespread on Ndumu Hill, not common.
Woodland.
D. viride (L.) Moench 476 Ndumu Hill. Woodland. Frequent but scattered.
Browsed by antelope.
. ‘ i dland.
1086A Ledebouria apertiflora (Bak.) Jessop 1811 Bulb. Ndumu Hill. Woo
L. revoluta ER 1812 Bulb. E slope Ndumu Hill in Acacia nigrescens
Thicket.
7272
1089
1090
1113
(28)
1110
(31)
1167
1168
1189
Journal of South African Botany
Ornithogalum longibracteatum Jacq. 535 Uluk. 162 Nd. Hill Bulb. Uluk-
ondo; Ndumu Hill. On Ndumu Hill, plants are much smaller in all respects than
Ulukondo specimens. In Sand Forest the bulbs grow above the ground. Sap from
the bulb stings fiercely. Ndumu Hill bulbs are very much smaller and are deeply
buried. Bushpigs and antelope eat entire plant (both forms).
Drimiopsis maculata Lindl. 35 T&W Bulb. Ndumu Hill in thickets, in wood-
land.
Asparagus acocksii Jessop 1137 Climber. Mahemane Thicket. Sickly sweet
scent.
A. africanus Lam. 72 Climber. Ndumu Hill. Woodland. Very attractive scent.
A. buchananti Bak. 1348 Climber. Mahemane Thicket.
A. densiflorus (Kunth) Jessop 1145 Herb. Mahemane Thicket. Swollen tubers
on roots.
A. falcatus L. var. falcatus 847 Climber. Ulukondo; Mahemane. Common in
thicket and Sand Forest. Sweetly scented. Browsed by antelope. Monkeys eat
shoots and leaves.
A. falcatus L. var. ternifolius (Bak.) Jessop 863 Climber. Widespread and
common in thicket and woodland. Strongly scented. Browsed.
A. macowanii Bak. var. zuluensis (N.E.Br.) Jessop 861 Shrub. Ulukondo,
Sand Forest. New spike produced each year which develops into a shrub—dies
back after a couple of years.
A. minutiflorus (Kunth) Bak. 1486 Herb. E slope of Ndumu Hill. Acacia
nigrescens Woodland. Scented.
A. saundersiae Bak. 1152 Climber. Mahemane Thicket.
A. setaceus (Kunth) Jessop 1316 Climber. Acacia tortilis Woodland in west.
A. subulatus Thunb. 1138 Climber. Mahemane, locally common in thicket.
A. virgatus Bak. 873 Herb. Ndumu Hill; Mahemane. Woodland and thicket.
Monkeys eat whole plant.
AGAVACEAE
Sansevieria desertii N.E.Br. 1437 Bulb. Mahemane. In thicket, forming large
communities. Flowers open in evenings, droop by morning. Scented. Do not
flower every year. Spike on tip of leaf raises sore bump on human skin.
Occasionally eaten by rhino and antelope.
S. grandis Hook.f. var. zuluensis N.E.Br. 1816 Bulb. Ulukondo. Locally
common in Sand Forest. Flowers caramel-toffee scented, open in evening.
Browsed by nyala.
S. hyacinthoides (L.) Druce 817 Bulb. Widespread, common. Forms large
communities especially in disturbed areas in thicket and woodland. Fibre from
leaves used for binding twine in manufacture of baskets. Browsed by rhino and
antelope. Mole rats eat the rootstock.
AMARYLLIDACEAE
Haemanthus multiflorus Mart. 660 Bulb. Widespread on Ndumu Hill; also in
west, in woodland and thicket. Plants usually found in shade. Leaves appear with
flowers. Browsed by antelope. Bushpig eat entire plant.
Boophane disticha (L.f.) Herb. 982 Bulb. Mkonjane. Acacia nigrescens
Woodland on rocky ground.
Crinum acaule Bak. 106 Bulb. Ndumu Hill, woodland, common. Flowers
open late afternoon and close at about 08h00 next day. Ovary buried in sand with
fruits developing just below or at surface. Leaves continually browsed, and
flowers too. They seldom survive the night. Very heavy, lovely scent. Flowers
1314
(38)
1422a
1568
1648
Checklist of Plants in Ndumu Game Reserve 23
normally open with storm or rain and plant can bloom a number of times a year.
pears grow continuously unless browsed. Antelope browse. Bushpigs eat the
ulb.
C. delagoense Verdoorn 1502 Bulb. Ndumu Hill, infrequent. Woodland.
C. macowanii Bak. 707 Bulb. Frequent on margins of Pongola Floodplain.
Scented. Buds and stem held upwards. As flowers open, stem rests on ground.
Flowers open upwards. Some leaves 2 m long and 100 mm wide in deep shade.
C. paludosum Verdoorn 282 Bulb. Frequent in west in seasonal pools and on
drainage lines. Faintly scented.
HYPOXIDACEAE
Hypoxis angustifolia Lam. 105 Bulb. E slope Ndumu Hill.
H. rooperi S. Moore 662 Bulb. Ndumu Hill. Woodland. Flowers open early
and close at midday.
DIOSCOREACEAE
Dioscorea cotinifolia Kunth 694 Climber. Mahemane, common. Thicket and
Sand Forest. Pretty pink galled fruits found quite frequently.
D. sylvatica (Kunth) Eckl. 539 KLT Climber. In thicket.
IRIDACEAE
Dietes flavida Oberm. 657 Herb. Mkonjane. In thicket on rhyolite outcrops.
Flowers open late morning and close at dusk. Browsed by antelope.
Gladiolus dalenii van Geel 1652 Bulb. Infrequent on Ndumu Hill, in thickets
in woodland, (perhaps for protection from bushpigs which eat the bulbs with
relish).
G. sp. 1611 Bulb. Banks of Pongola river. More material needed. Flowers
large white/pink.
Lapeirousia grandiflora Bak. 796 Bulb. Ndumu Hill. Infrequent, usually
found on edge of thickets. First day, bud opens slowly, second day open fully,
pollen visible—third day pollen lost, flower closes up slowly in afternoon. Drops
next day. Orange seeds germinate readily. Bushpig eat the corms.
ORCHIDACEAE oe
Bonatea boltonii (Harv.) Bolus 1398 Bulb. East of Pongola Floodplain in
thicket in open woodland.
B. saundersiae Harv. 633 Bulb. Mahemane Thicket. Leaves usually dead at
time of flowering. Spur unfurls weeks before flower opens. Browsed by antelope.
Ansellia gigantea Reichb. f. var. nilotica Summerh. 654 Epiphyte. Mahemane;
Ulukondo; Engabateni; east of Pongola Floodplain. Fairly common in Sand Forest
and thicket. Huge clumps of roots and pseudobulbs. Scented. Flowers last a long
time (weeks). Known by Tongas as the ‘monkey’s sugar cane because the
monkeys eat the plant. Large clumps were placed in trees in the camp in the early
1960’s.
Eulophia hereroensis Schltr. 96 Bulb. Ulukondo. In Sand Forest and thickets
in woodland. Bushpig eat the rootstock.
E. leachii Gace Ze Hall 304 Bulb. Mahemane Thicket. Flowers appear
with new leaves. Browsed by antelope.
E. petersii Reichb.f. 175 Bulb. Mahemane; Mkonjane. In thicket and on
thyolite outcrops in west. Grow in large clumps—inflorescences will grow to a
height of 2 m and more (when in shade) to reach the light.
24
1835a
1837
Journal of South African Botany
Cyrtorchis praetermissa Summerh. 1202 Epiphyte. Ulukondo, Sand Forest.
Not common, found in valley-mist areas.
Mystacidium capense (L.f.) Schlechter 852 Epiphyte. Ulukondo Sand Forest.
In valley-mist belt, seldom found below 2 m. Scented at night.
1837a Microcoelia exilis Lindl. 353 Epiphyte. Ulukondo Sand Forest and on trees
adjacent to Inyamiti and Banzi pans.
DICOTYLEDONEAE
(43) ULMACEAE
1902 Trema orientalis (L.) Bl. 839 Tree. Usutu Floodplain and in disturbed areas in
(44)
1961
(46)
1980
(48)
2074
2093
(49)
2118
(50)
2131
Riverine Forest. Birds eat fruits. Browsed by antelope. Young plants in shade of
forest can have very large leaves.
MORACEAE
Ficus capreifolia Del. 398 Shrub. Floodplains and especially the Pongola river
banks. Browsed. Used for making Fonya fishing baskets and the leaf is used as
sandpaper by the Tongas when carving household utensils.
F. sycomorus L. 823 Tree. Floodplains and Riverine Forest. Trees can be
found fruiting at almost any time of the year. The fruits can be very sweet. The
Tongas mash them up with water when food is scarce at the end of winter. Birds,
monkeys, bushbabies, antelope, bushpig and people eat the figs (umKhiwane).
The leaves are browsed by antelope. Ripe fruits are used as bait for catching
barbel.
URTICACEAE
Laportea peduncularis (Wedd.) Chew 396 Shrub. Usutu Riverine Forest.
LORANTHACEAE—the family is attractive to sunbirds.
Loranthus bolusii Sprague 1675 Parasite. Ndumu Hill. Thicket in woodland
and in pan-edge community on Acacia grandicornuta.
L. dregei Eckl. & Zeyher 872 Parasite. Pongola Floodplain, Ndumu Hill. On
Trichilia emetica.
L. kalachariensis Schinz 1364 Parasite. Ndumu Hill. On Acacia burkei.
L. kraussianus Meisn. 812 Parasite. Mahemane Thicket, floodplains (in
thickets).
L. minor (Harv.) Sprague 1676 Parasite. S bank Inyamiti in thicket. On
Maytenus heterophylla. Faintly scented.
L. ngamicus Sprague 801 Parasite. Mahemane Thicket. On Acacia tortilis.
L. oleaefolius Cham. & Schlechtd. 702 JHR Parasite. Ndumu Hill. on Acacia
burkei.
L. quinquenervis Hochst. 1436 Parasite. Mahemane Thicket.
Viscum spragueanum Burtt Davy 1481 Parasite. Ndumu Hill. On Strychnos
spinosa.
V. verrucosum Harv. 1363 Parasite. Mahemane Thicket on Acacia grandicor-
nuta. Forms large masses 1 m in diameter.
SANTALACEAE
Thesium sp. 1660 Herb. Ndumu Hill. Woodland.
Thesium sp. 139 Herb. Mahemane Thicket. Common.
OLACACEAE
Olax dissitiflora Oliv. 1388 Tree. Mkonjane. Uncommon in Acacia nigrescens
Woodland.
2136
(52)
2201
2204
(53)
2223
2255
2257
(54)
2292
2293
2299
2309
2311
2312
2314
2324
2328
Checklist of Plants in Ndumu Game Reserve 25
Ximenia caffra Sond. var. natalensis Sond. 1173 Shrub. Mahemane Thicket.
Fruits eaten by monkeys, birds and man (amaThunduluka). No. 715, from Ndumu
Hill where it is common, differs in leaf colour and size but good flowering
material is needed for correct determination.
POLYGONACEAE
Polygonum aviculare L. 1035 Herb. Edge of Banzi pan.
P. plebeium R.Br. 528 KLT Herb. Dried mudflats.
P. pulchrum Blume 200 Herb. Pongola Floodplain. Common.
P. senegalense Meisn. forma albotomentosum R. Grah. 1358 Herb. Pongola
Floodplain. Common in scattered patches.
Oxygonum dregeanum Meisn. var. dregeanum 1809 Herb. Ndumu Hill. Com-
mon in woodland.
CHENOPODIACEAE
Chenopodium album L. 297 Herb. Ndumu Hill. Very common in woodland
especially on disturbed ground. Unpleasant scent.
C. ambrosioides L. 491 Herb. Usutu River banks. Common.
Arthrocnemum natalense (Bunge ex Ung-Sternb.) Moss var. natalense 460
KLT Inyamiti pan, mudflats. Uncommon.
Salicornia pachystachya Bunge ex Ung-Sternb. 888 Succulent herb. Inyamiti
mudflats and stream. Browsed by antelope.
AMARANTHACEAE
Celosia trigyna L. 1649 Herb. Ulukondo, locally common. Sand Forest.
Hermbstaedtia odorata (Burch.) T. Cooke 149 Herb. Widespread especially in
disturbed areas. Thorny fruit.
Amaranthus thunbergii Mog. 1508 Herb. Common on Ndumu Hill; margins
Pongola Floodplain—on disturbed soil.
Cyphocarpa angustifolia Lopr. 1198 Herb. Red Cliffs. Acacia tortilis Thicket.
Bunguzane. Browsed by nyala.
Centema subfusca (Moq.) Lopr. 19 TBO Acacia xanthophloea Forest, east of
Banzi pan.
Cyathula spathulifolia Lopr. 846 Herb. Ulukondo. Common in Sand Forest.
Burr fruit.
Pupalia atropurpurea Mog. 719 Scandent herb. Widespread, common espe-
cially in thickets. Clinging burr fruit.
Nothosaerva brachiata (L.) Wight 1560 Herb. Mahemane pools (Ist record
for Natal).
Psilotrichum africanum Oliv. 733 Herb. Ulukondo. On edge of Sand Forest.
Achyranthes aspera L. 465 Herb. Ndumu Hill, in thicket; Riverine Forest.
C ‘ . “ys
ea (L.) All. 1310 Herb. Mahemane Thicket; Acacia tortilis Woodland.
2328a Achyropsis leptostachya Hook.f. 1195 Herb. Margins of floodplains.
26 Journal of South African Botany
2335 Alternanthera pungens H.B.K. 1507 Prostrate herb. Widespread especially on
disturbed ground. ‘Paper thorns’.
A. sessilis (L.) DC. 421 KLT ‘‘Common emergent aquatic, usually in water up
to 6 m deep”’ (KLT)
2338 Gomphrena celosioides Mart. 1595 Herb. Mahemane, widespread. On edges
of seasonal pools.
(55) NYCTAGINACEAE
2347a Commicarpus africanus (Lour.) Dandy 666 Common on south boundary fence-
line in the west. Herb.
2349 Boerhavia diffusa L. var. hirsuta Heim. ex descr. 415 Herb. Common on
Ndumu Hill. Woodland. Fruits stick to clothing.
2354 Pisonia aculeata L. 1418 Climber. Common in Riverine Forest. No flowers or
fruits have been collected at Ndumu.
(57) MOLLUGINACEAE
2376 Limeum fenestratum (Fenzl) Heimerl 754 Herb. Ndumu Hill, woodland. Com-
mon.
L. viscosum (J. Gay) Fenzl subsp. viscosum var. glomeratum (Eckl. & Zeyher)
Freidr. 1238 Herb. Ndumu Hill, drainage lines.
2382 Gisekia africana (Lour.) O. Kuntze var. africana 150 Prostrate herb. Edge of
Pongola Floodplain. Common.
G. africana (Lour.) O. Kuntze var. cymosa Adamson 296 Herb. Ndumu Hill.
Common. Woodland.
Mollugo nudicaulis Lam. 1552 Herb. Seasonal pools. Common (1st record for
Natal).
2388 Glinus oppositifolius A.DC. 364 Herb. Banzi pan margins.
2390 Hypertelis bowkeriana Sond. ex descr. 1607 Herb. Bunguzane. Thicket, on
disturbed ground.
2393a Corbichonia decumbens (Forsk.) Exell 360 Herb. East slope Ndumu Hill;
Acacia nigrescens Woodland along base of Ndumu Hill. Flowers open midday to
sunset and re-open next day.
(58) AIZOACEAE
2395a Zaleya pentandra (L.) Jeffrey 94 Herb. Common in Mahemane Thicket.
2403 Tetragonia expansa Thunb. 1465 Herb. Pongola Floodplain margin (intro-
duced).
(59) MESEMBRY ANTHEMACEAE
2405 Aptenia cordifolia (L.f.) N.E.Br. 13 TBO Herb. Thicket.
2405 Delosperma lebomboense (L. Bol.) Lavis 1266 Herb. Common in Mahemane
Thicket.
(60) PORTULACACEAE
2406 Talinum caffrum (Thunb.) Eckl. & Zeyher 303 Herb. Widespread in thicket
and woodland. Flowers open afternoon to night. Plants die back in winter,
re-shoot from carrot-like rootstock. Bushpig eat the rootstock.
Checklist of Plants in Ndumu Game Reserve 27
Talinum portulacifolium (Forsk.) Aschers. ex Schweinf. 201 Succulent herb.
Ulukondo, locally common on edge of Sand Forest and thicket. Flowers open at
about 15h30 and close in late afternoon. Blooms all summer and dies back in
winter to re-shoot from same rootstock—swollen roots, not as well developed as
the previous species. Browsed by antelope.
2419 Portulacaria afra Jacq. 842 Succulent shrub. Widespread, particularly at old
kraal sites. Occasionally browsed.
2421 Portulaca pilosa L. 1277 Herb. Mahemane Thicket.
P. quadrifida L. 1636 Herb. Mahemane Thicket.
(61) BASELLACEAE
2424 Basella paniculata Volkens 4891 EJM_ Climber. Mahemane Thicket.
(63) ILLECEBRACEAE
2465 Pollichia campestris Ait. 295 Herb. Ndumu Hill. Common in woodland on
Hutton red sands.
(64) NYMPHAEACEAE
2513 Nymphaea capensis Thunb. 585 Aquatic. Widespread in pools and pans.
Flowers remain open for some days. Heavy scent, stronger at night. Hippo eat
tubers, monkeys eat exposed roots on mudflats. Tubers collected by Tongas to
take home and cook (amaZibo).
N. lotus L. 765 Aquatic. Banzi pan, Inyamiti stream. Flowers open at night,
close mid-morning. Tubers eaten by man and beast—see above (amaHlolwane).
(65) CERATOPHYLLACEAE
2516 Ceratophyllum demersum L. 1758 EJM ‘‘Submerged aquatic in open fresh
water’ (EJM).
(66) RANUNCULACEAE
2546 Ranunculus multifidus Forsk. 165 Herb. Floodplains. Common.
(67) MENISPERMACEAE
2570 Cocculus hirsutus (L.) Diels 545 Climber. Widespread, common especially in
woodland. Browsed.
2574 Cissampelos hirta Klotzsch 2016 CJW Climber. Ndumu Hill. Common.
Woodland. Browsed.
C. torulosa E. Mey. ex Harv. 86 Climber. Ndumu Hill. Woodland.
2643 Epinetrum delagoense (N.E.Br.) Diels 529 Climber. Ndumu Hill, woodland.
Forms dense communities. Fruits eaten by Tongas (inGantaganta).
(68) ANNONACEAE
i i Forest.
2673 Uvaria caffra E. Mey. ex Sond. 783 Climber. Ulukondo. Sand
U. lucida Benth. subsp. virens (N.E.Br.) Verde. 1003 KLT Climber. Engaba-
teni. Sand Forest.
2696 Monanthotaxis caffra (Sond.) Verdc. 450 Climber. Widespread, common in
woodland and thicket. Browsed by antelope. Birds eat the fruits.
2733 Monodora junodii Engl. & Diels 1412 Tree. Ulukondo, Sand pore taas
occasional along the Pongola Floodplain. Fruits eaten by Tongas (umKhotshi).
28
(70)
2813
2825
(71)
2852
(73)
2883
2965a
(74)
3082
3099
3101
3106
3109
3112
Sil M3)
Journal of South African Botany
LAURACEAE
Cryptocarya woodti Engl. 847 KLT Pongola Riverine Forest.
Cassytha filiformis L. 1622 Climber. Ndumu Hill. Frequent in woodland.
PAPAVERACEAE
Argemone mexicana L. 75 Herb. Usutu Floodplain (introduced).
BRASSICACEAE (nom. alt. Cruciferae)
Lepidium africanum (Burm.f.) DC. 1439 Herb. Ndumu Hill. Common on
disturbed ground.
Rorippa madagascariensis (DC.) Hara 1464 Herb. Common on Pongola
Floodplain.
CAPPARACEAE
Cleome angustifolia Forsk. subsp. diandra (Burch.) Kers. 391 Herb. Ndumu
Hill. Frequent in woodland. Flowers close at midday.
C. monophylla L. 370 Herb. Pongola Floodplain margins. Common. Flowers
close midday.
Cladostemon kirkii (Oliv.) Pax & Gilg 29 Tree. Ndumu Hill, towards Bungu-
zane and S bank Inyamiti; Ulukondo. Flowers open green, turning white then
yellow, petals veined in green. Scented. Flowers appear with new leaves.
Browsed by antelope. Roots spread out horizontally from tree. Small piece of root
will grow into a tree. Afflicted by a borer. Ripe fruits stink.
Capparis brassii DC. 108 Climber. Mahemane, widespread in thicket.
Browsed by antelope.
C. fascicularis DC. var. fascicularis 628 Climber. Mahemane Thicket. Pleas-
antly scented flowers.
C. sepiaria L. var. citrifolia (Lam.) Toelken 1267 Scandent shrub. Mahemane
Thicket. Particularly common in Acacia tortilis Thicket.
C. tomentosa Lam. 85 Climber. Common, especially on floodplains. Wide-
spread. Flowers wilted by early afternoon. Fruits eaten by monkeys and man
(umQokolo, uKokwane). Browsed by nyala.
Boscia albitrunca (Burch.) Gilg & Ben. 1197 Tree. Widespread. Much
browsed. Insects and birds enjoy the flowers and fruits.
B. foetida Schinz subsp. rehmanniana (Pest.) Toelken 833 Shrub. Mahemane,
Ulukondo. Thicket and Sand Forest. Sickly sweet scent. Browsed by antelope.
Cadaba natalensis Sond. 826 Shrub. Widespread, particularly in thicket and
woodland. Browsed by antelope.
Maerua angolensis DC. 721 Tree. Mahemane. Thicket and woodland.
Browsed by antelope.
M. edulis (Gilg & Ben.) De Wolf 79 Shrub. Mahemane Thicket. Common on
roadsides.
M. juncea Pax subsp. crustata (Wild) Wild 219 Climber. Mahemane. Thicket
and woodland. Flowers open for a day. Faintly scented. Fruits orange when ripe.
Browsed by nyala.
M. rosmarinoides (Sond.) Gilg & Ben. 581 KLT Shrub. Bunguzane.
Thylachium africanum Lour. 80 Shrub. Widespread; common in woodland,
thicket and Sand Forest. Flowers close after a day. Much browsed by all antelope.
Flowers white becoming pink. Fruits eaten by Tongas (isiKonke).
(79)
3164
3166
3168
(87)
3353
(90)
3443
3446
Checklist of Plants in Ndumu Game Reserve 29
CRASSULACEAE
Cotyledon wickensii Harv. 520 Succulent. Mahemane Thicket.
Kalanchoe rotundifolia Haw. 14 Succulent. Widespread. Common, especially
in thicket. Browsed.
K. rotundifolia (Haw.) Harv. var. peltata Raymond-Ham. 307 Succulent:
Ndumu Hill. Uncommon. Woodland. Browsed.
K. sp. 580 Succulent. Mkonjane; E slope Ndumu Hill. Acacia nigrescens
Woodland. Plant takes 2 to 3 years to mature. Dies back after flowering but
re-shoots from base.
Crassula portulacea Lam. 16 Succulent. Mahemane Thicket and Sand Forest.
C. transvaalensis Kuntze 288 Herb. Ndumu Hill. Infrequent. Woodland.
C. sp. 468 Herb. Inyamiti pan margins; Ulukondo; Mahemane. Common in
thicket and Sand Forest.
ROSACEAE
Rubus rigidus Smith 171 Climber. Pongola Floodplain. Fruits eaten by birds
and Tongas (uGagane).
LEGUMINOSAE
Albizia adianthifolia (Schumach) W. F. Wight 1414 Tree. E of Pongola
Floodplain.
A. anthelmintica (A. Rich.) A. Brongn. 649 Tree. Dominant in Mahemane;
elsewhere in thicket. Conspicuous in August/September when in full flower
without leaves. Lovely scent. Browsed by antelope.
A. forbesii Benth. 1403 Tree. Ulukondo. Sand Forest. Browsed.
A. petersiana (Bolle) Oliv. subsp. evansii (Burtt Davy) Brenan 980 Tree.
Ndumu Hill. Woodland; widespread. Browsed by antelope. Trees have coppery
spring foliage.
A. versicolor Welw. ex Oliv. 710 Tree. Ndumu Hill. Woodland. Flowers
appear on new shoots. Attractive scent. Parrots eat seeds out of pods.
Acacia albida Del. 3151 CJW Tree. Usutu Floodplain. Uncommon. Some
very large specimens. ;
A. borleae Burtt Davy 825 KLT Tree. Isolated patch on road from main gate
to Inyamiti.
A. burkei Benth. 527 Tree. Ndumu Hill. Woodland, very common. Flowers
unpleasantly scented. Favoured for bird nests perhaps because of the protective
vicious thorns. Gum eaten by monkeys and Tongas. Browsed by antelope.
A. caffra (Thunb.) Willd. 655 JHR Tree. Mkonjane. Not common, in Acacia
nigrescens Woodland. '
A. gerrardii Benth. var. gerrardii 1382 Tree. Frequent in western area, scat-
tered elsewhere in woodland and thicket. ; ;
A. grandicornuta Gerstner 1177 Tree. Mahemane. Dominant. Widespread in
thicket. Browsed.
A. kraussiana Meisn. ex Benth. 1401 Climber. E of Pongola Floodplain on
edges of Sand Forest. ;
A. luederitzii Engl. var. retinens (Sim) Ross & Brenan
Thicket and woodland near drainage lines in west. ii
A. nigrescens Oliv. 897 KLT Tree. Common in the western area; Ndumu Hi
slopes. Browsed. .
A. nilotica (L.) Willd. ex Del. subsp. kraussiana (Benth.) Brenan 1270 Tree.
Ndumu Hill. Widespread, dominant in parts of west. Beautiful scent. Monkeys eat
flowers; monkeys and people eat the gum. Browsed.
1125 Tree. Mahemane.
30
Journal of South African Botany
Acacia robusta Burch. subsp. clavigera (E. Mey.) Brenan 790 KLT Tree.
Margins of pans, floodplains.
A. schweinfurthii Brenan & Exell var. schweinfurthii 1400 Climber. Riverine
Forests. Leaves close in dark or when picked.
A. senegal (L.) Willd. var. rostrata Brenan 1158 Shrub. Widespread but not
common. Browsed.
A. tortilis (Forsk.) Hayne subsp. heteracantha (Burch.) Brenan 595 Tree.
Widespread, common—forms pure stands. Browsed. Dried seed pods eaten by
antelope.
A. xanthophloea Benth. 696 JHR_ Tree. Floodplains, forms pure stands at
Shokwe and Banzi. Monkeys eat leaves, buds, flowers, seeds.
3449 Mimosa pigra L. 166 Shrub. Pongola Floodplain.
3452 Dichrostachys cinerea (L.) Wight & Arn. Two different forms found on Ndumu
Hill, woodland; KLT 785 (large leaf) becoming tree-like with black striated bark.
ESP 964 (small leaf) remains a many stemmed shrub. Browsed. Nyala eat seed
pods. Monkeys eat seeds.
3460 Newtonia hildebrandtii (Vatke) Torre var. hildebrandtii 1111 Tree. Dominant
in Ulukondo. Sand Forest. Pink-winged seeds litter the forest floor beautifully.
3506 Schotia brachypetala Sond. 12 Tree. Ndumu Hill, edge of floodplains. Many
insects and birds, particularly sunbirds, attracted to flowers. Flowers appear with
the new leaves. Leaves browsed. Monkeys eat flower buds. Bark of tree used to
dye Tonga fish nets.
S. capitata Bolle 1093 Tree. Mahemane, in thicket. Browsed. Sunbirds parti-
cularly attracted to flowers.
3509 Afzelia quanzensis Welw. 1438 Tree. East of Pongola Floodplain. One young
specimen in Acacia xanthophloea Forest at Banzi, perhaps seed brought down by
floods (from Lebombo mountains).
3528 Bauhinia galpinii N.E.Br. 1655 Shrub. Fontana pan margins, Paphekulu
stream. Scandent shrub. Planted in camp and crocodile farm gardens.
3530 Dialium schlechteri Harms 156 Tree. Ndumu Hill. Woodland, Sand Forest.
Heavy rather unpleasant scent. People, monkeys and birds eat the fruits
(umThiba).
3536 Cassia bicapsularis L. 641 Scandent shrub. Usutu Riverine Forest.
C. italica (Mill.) Lam. ex F. W. Anders subsp. arachoides (Burch.) Brenan
92 Sub-woody plant, Ndumu Hill. Woodland. Frequently found on roadsides.
C. mimosoides L. sens. lat. 716 Herb. Ndumu Hill. Woodland. Frequent.
C. occidentalis L. 405 Shrub. Usutu Floodplain.
C. petersiana Bolle 499 Shrub. Common on Ndumu Hill. Woodland. Sweetly
scented. Monkeys eat green pods. Birds eat seeds. Tongas eat seeds (umNem-
benembe). Browsed. Forms thickets on the hill and spreads quickly giving cause
for concern.
C. siamea Lam. 1515 CJW_ Tree. At old kraal site, Usutu Floodplain.
3551 Parkinsonia aculeata L. JEWD_ Tree. Catuane crossing. Riverine Forest (intro-
duced).
3561 Peltophorum africanum Sond. Tree. Ndumu Hill. Woodland.
3607a Bolusanthus speciosus (Bolus) Harms 19 Tree. Widespread, in woodland and
on edges of Usutu Floodplain. Flowers appear with new leaves. Faintly scented.
3669
3688
3702
3718
3719
3720a
3747
3792
3802
3804
3810
Checklist of Plants in Ndumu Game Reserve 31
Small worms eat colour off flowers and the leaves. Flower buds eaten by vervet
monkeys.
Crotalaria near C. burkeana Benth. 429 Herb. S slope Ndumu Hill. Wood-
land.
C. monteiroi Taub. ex Bak.f. 203 Shrub. Ndumu Hill; Usutu Floodplain.
Browsed.
C. natalensis Bak.f. TBO Herb. SW corner of reserve.
C. pallida Ait. var. pallida 534 Herb. Pongola Floodplain.
m aba othe Perr. ex DC. 416 Herb. Ndumu Hill. Frequent. Woodland, in
shade.
Medicago aschersoniana Urb. 1693 Herb. Ulukondo, Sand Forest. Browsed.
Indigofera arrecta Hochst. ex A. Rich. 1281 Herb. Usutu Floodplain. Mahe-
mane clearing.
I. charleriana Schinz var. charleriana 1442 Herb. Pongola Floodplain margin.
I. charleriana Schinz var. sessilis (Chiov.) Gillett 1589 Herb. Pongola Flood-
plain margin.
I. costata Guill. & Perr. subsp. macra (E. Mey.) Gillett 1252 Herb. Wide-
spread.
I. delagoensis Bak.f. ex Gillett 1554 Herb. Ndumu Hill. Common. Woodland.
I. filipes Benth. ex Harv. 1590 Herb. Ndumu Hill drainage line.
I. schimperi Jaub. & Spach var. schimperi 664 Herb. Mkonjane. Common on
disturbed areas.
I. spicata Forsk. 169 Herb. Pongola Floodplain. Frequent.
I. vicioides Jaub. & Spach var. rogersii (R. E. Fries) Gillett 1565 Herb.
Mahemane pools.
Tephrosia forbesii Bak. subsp. inhacensis Brummitt 363 Herb. Ndumu Hill.
Woodland. Flowers open mid-afternoon. Browsed.
T. longipes Meisn. subsp. longipes 753 Herb. Ndumu Hill; Mkonjane. Wood-
land. Browsed.
T. semiglabra Sond. 1582 Herb. Ndumu Hill. Common. Woodland.
T. sp. 1259 Herb in Acacia tortilis Woodland.
T. sp. 1285 Herb. Ndumu Hill, woodland.
T. sp. 308 Herb. Ndumu Hill, woodland.
Mundulea sericea (Willd.) A. Chev. 674 Shrub. Ndumu Hill; Mkonjane.
Woodland. Browsed by antelope.
Craibea zimmermannii (Harms) Harms ex Dunn 696 Tree. Ulukondo, Sand
Forest. Beautiful when in flower, scented.
Sesbania sesban (L.) Merrill subsp. sesban var. nubica Chiov. Pongola Flood-
plain. There appear to be distinct varieties:
541—Perennial tree, with many leaves.
170—Annual shrub, with few leaves.
Ormocarpum trichocarpum (Taub.) Engl. 427 Small tree. Ndumu Hill. Wood-
land. Browsed.
Stylosanthes fruticosa (Retz.) Alston 1236 Herb. Ndumu Hill. Woodland.
Zornia capensis Pers. 1692 Herb. Ndumu Hill. Woodland.
Alysicarpus glumaceus (Vahl) DC. non (Roth) Schindl. 726 Herb. Pongola
Floodplain margins.
32
3821
3828
3834
3856
3864
3870
3891
3892
3897
3898
3905
3910
3910d
Journal of South African Botany
Dalbergia armata E. Mey. Climber. Riverine Forest.
D. obovata E. Mey. 1214 Tree. Ndumu Hill. Infrequent. Woodland.
D. sp. c.f. D. nitidula Welw. ex Bak. 1659 Scandent shrub. Common in
Ulukondo. Sand Forest. Collected in flower and pod in February, 1972—the first
such flowering and fruiting observed in four years. Flowers appear without leaves.
Scented. Browsed.
Pterocarpus angolensis DC. TPD Tree. Usutu river bank. Rare.
P. rotundifolius (Sond.) Druce subsp. rotundifolius 1032 KLT Shrub. Mkon-
jane.
Lonchocarpus capassa Rolfe 1286 Tree. Scattered in western areas.
Abrus laevigatus E. Mey. 243 Climber. Mkonjane. Thickets in Acacia nigres-
cens Woodland and on rhyolite outcrops.
A. precatorius L. subsp. africanus Verde. 572 Climber. Floodplains and
Ndumu Hill. Seeds used for decoration by Tongas—in musical instruments and
clay pots.
Glycine wightii (R. Grah. ex Wight & Arn.) Verde. 1315 Climber. Usutu
Floodplain. Acacia tortilis Woodland. In shade of continuous canopy of forest
patches.
Erythrina humeana Spreng. 815 Shrub with large swollen underground root.
Ndumu Hill, woodland. Usually protected in thickets from browse, fire.
E. lysistemon Hutch. Tree. Usutu Floodplain at old kraal sites.
Canavalia virosa (Roxb.) Wight & Arn. 1362 Climber. Pongola Riverine
Forest.
Cajanus cajan (L.) Millsp. 1347 Herb. Usutu Floodplain.
Rhyncosia minima (L.) DC. var. minima 720 Climber. Common on Ndumu
Hill. Flowers closed in afternoon.
R. minima (L.) DC. var. prostrata (Harv.) Meikle 1615 Climber. S boundary,
in west.
R. totta (Thunb.) DC. var. totta 722 Climber. Pongola Floodplain.
Eriosema psoraleoides (Lam.) G. Don 1342 Shrub. Floodplains. Forms occa-
sional communities. Tongas collect seeds in large quantities, take home and cook
mixed with mealie meal, etc. (uTongolo).
Vigna luteola (Jacq.) Benth. 358 Climber. Banzi pan, on floating rafts of
vegetation.
V. unguiculata (L.) Walp. var. protracta (E. Mey.) Verde. 252 Climber.
Ndumu Hill. Common in woodland. Flowers close before noon. Browsed. Tubers
on roots eaten by Tongas.
V. vexillata (L.) Rich. sens. lat. 173 Climber. Pongola Floodplain.
Dolichos trilobus L. subsp. transvaalicus Verdc. 600 Climber. Usutu Flood-
plain.
Macrotyloma axillare (E. Mey.) Verde. 1654 Climber. Ndumu Hill. On the
26/2/72 (ESP) and 25/3/55 (ICP) this creeper covered the grasses and shrubs of
Ndumu Hill to the point of ‘suffocation’—bleaching of grasses. Browsed, or
eaten with grasses by nyala.
M. maranguense (Taub.) Verde. 1824 Climber. Ndumu Hill. Tubers on roots
dug up and eaten by Tongas, raw and cooked. Bushpigs also dig up to eat
(isiKhondle).
Checklist of Plants in Ndumu Game Reserve 33
(91) GERANIACEAE
3928 Pelargonium sp. 2564 CJW Herb. Ndumu Hill.
(92) OXALIDACEAE
3936 Oxalis corniculata L. 709 Herb. Ndumu camp. (Weed) spread by birds which
eat seeds.
O. semiloba Sond. subsp. semiloba 1292 Herb. Scattered communities on
Ndumu Hill and in the west, on roadsides. Flowers close mid-afternoon.
(94) ERYTHROX YLACEAE
3956 Erythroxylon emarginatum Thonn. 1677 Tree. Ulukondo. Sand Forest.
(95) ZYGOPHYLLACEAE
3978 Tribulus terrestris L. 1602 Herb. Ndumu Hill, Pongola Floodplain. On dis-
turbed ground. ‘Devil Thorn.’
(96) BALANITACEAE
3980 Balanites maughamii Sprague 591 Tree. Mahemane Thicket. Common.
Browsed. Nyala and monkeys eat fruits.
B. pedicellaris Mildr. & Schltr. 311 Tree. Mahemane. Common in thicket.
(97) RUTACEAE
4076 Vepris carringtoniana Mendonca 685 Shrub. Mahemane near main gate and S
boundary fence. Only known locality in Natal.
4078 Toddaliopsis bremekampii Verdoorn 860 Shrub. Ulukondo. Frequent in Sand
Forest.
4091 Clausena anisata (Willd.) Hook.f. ex Benth. 809 Shrub. Widespread in
thicket. Heavy unpleasant scent when leaf or stem is broken. Browsed.
(98) BURSERACEAE
4151 Commiphora neglecta Verdoorn 508 Tree. Ndumu Hill—frequently in large
communities in woodland, forming thickets. Browsed. Many species of bird eat
the fruits. Tree trunks cut and dried to make floats for fishing nets. Swollen roots
dug up and eaten by Tongas and, on a large scale, by bushpigs (umNyela).
C. glandulosa Schinz 723 Tree. E slope Ndumu Hill in Acacia nigrescens
Woodland. Bunguzane (1st record for Natal). '
C. pyracanthoides Engl. subsp. pyracanthoides 78 Shrub. Red Cliffs; Ndumu
Hill; western area in woodland.
(99) PTAEROXYLACEAE
4157 Ptaeroxylon obliquum Eckl. & Zeyher 811 Tree. Mahemane. Common in
woodland and thicket. Browsed. Good fire wood.
(100) MELIACEAE ne Rone
4163 Entandophragma caudatum (Sprague) Sprague 2657 / Tree. One large
Specie eet of Bunguzane. No young specimens known in the wild. Yellow
autumn foliage conspicuous from a great distance (the camp, Ndumu Hill).
4171 Turraea obtusifolia Hochst. 384 Shrub. Ndumu Hill; Mkonjane. Woodland.
Browsed.
4175 Melia azedarach L. 878 Tree. Riverine Forests. Monkeys eat flowers and
berries (introduced).
34
Journal of South African Botany
4193 Ekebergia capensis Sparrm. 841 Tree. Usutu Floodplain in forest patches.
4195
(101)
4219
4220
(102)
4273
(103)
4283
(104)
4297
429
\o
4309
4325
4327
4343
Trichilia emetica Vahl. subsp. emetica 824 Tree. Widespread. Sweetly scented
flowers. Browsed. Fruits eaten by birds, antelope, monkeys (also eat flower buds)
and the Tongas who collect the seeds, soak them, squeeze them, cook and eat as
soup (umKhuhlu). The wood is used extensively in the manufacture of household
utensils by Tongas, and for making drums.
MALPIGHIACEAE
Sphedamnocarpus pruriens (A. Juss.) Szyszyl. 478 Climber. Ndumu Hill.
Woodland, and thicket.
Acridocarpus natalitius A. Juss. var. linearifolius Launert Scandent shrub.
Ulukondo. Sand Forest. Browsed.
POLYGALACEAE
Polygala amatymbica Eckl. & Zeyher 112 Herb. E slope Ndumu Hill.
Thicket.
P. senensis Klotzsch 1558 Herb. Bunguzane; Mahemane pools (1st record for
Natal).
P. sphenoptera Fresen 126 Herb. Ndumu Hill, woodland.
P. producta N.E.Br. 133a Herb. Margins of Pongola Floodplain. Flowers
close at night.
DICHAPETALACEAE
Dichapetalum cymosum (Hook.) Engl. 890 Herb. Very common on Ndumu
Hill. Apparently impervious to fire—is the first to shoot again after a burn. 1671
on Ndumu Hill above Ulukondo are very large communities of very hairy plants.
EUPHORBIACEAE
Securinega virosa (Roxb. ex Willd.) Pax & Hoffm. 1279 Shrub. Widespread.
Sweetly scented flowers, attract many insects. Birds feed on fruits. Browsed.
Phyllanthus asperulatus Hutch. 1605 (The status of this species is uncertain)
Herb. Mdipine. Acacia tortilis Woodland.
P. burchellii Muell. Arg. 1033 Herb. Ndumu Hill. Common in woodland.
Leaves close at night or on picking.
P. discoideus (Baill.) Muell. Arg. 942 KLY E of Pongola Floodplain.
Browsed.
P. flacourtioides Hutch. 1640 Tree. Pongola Riverine Forest.
P. maderaspatensis L. 724 Herb. Ndumu Hill. Very common.
P. reticulatus Poir. 838 Shrub. Floodplains. Browsed.
Drypetes arguta (Muell. Arg.) Hutch. 782 Tree. Ulukondo. Frequent in Sand
Forest.
Hymenocardia ulmoides Oliv. 1410 Tree. Ulukondo. Sand Forest; also occa-
sionally in woodland on Ndumu Hill. Used for making Fonya fishing baskets
(thrust baskets).
Antidesma venosum E. Mey. ex Tul. 573 Tree. Pongola Riverine Forest;
Ndumu Hill. Browsed. Monkeys, birds, Tongas eat fruits (umShongi).
Cleistanthus schlechteri (Pax) Hutch. 781 Tree. Ulukondo. Common in Sand
Forest.
Checklist of Plants in Ndumu Game Reserve 35
4345 Bridelia cathartica Bertol.f. subsp. cathartica 650 Shrub. Floodplains.
Browsed. Monkeys eat fruits.
4348 Croton gratissimus Burch. 865 Tree. Ulukondo. In Sand Forest. Used to make
furniture by Tongas.
C. menyhartii Pax 887 Shrub. Bunguzane; widespread and common. Used for
Fonya (thrust) fishing baskets.
C. pseudopulchellus Pax 868 Shrub. Ulukondo. Sand Forest.
C. steenkampiana Gerstner 348 Shrub. Ulukondo. Common on edges of Sand
Forest.
4361 Caperonia stuhlmannii Pax 348 Herb. Pongola Floodplain.
4407 Acalypha ecklonii Baill. 1433 Herb. Pongola Riverine Forest.
A. glabrata Thunb. 738 Shrub. Engabateni. Uncommon. Sand Forest. Used
for Fonya (thrust) fishing baskets.
A. indica L. 305 Herb. Ndumu Hill. Woodland; on disturbed ground.
A. petiolaris Hochst. 604 Shrub. Usutu Floodplain.
4416 Tragia incisifolia Prain 1597 Herb. Mkonjane. Acacia nigrescens Woodland.
T. sp. c.f. T. durbanensis O. Kuntze 772 Climber. Ndumu Hill. Frequent.
Plant raises stinging weals on skin. Browsed.
4422 Dalechampia galpinii Pax 679 Climber. Mkonjane. Acacia nigrescens Wood-
land in west.
4424 Ricinus communis L. 1813 Shrub. Floodplains (introduced. )
4433 Jatropha variifolia Pax 22 Shrub. Very common in Mahemane Thicket.
J. sp. c.f. J. curcas L. 466 Tree. Usutu Floodplain. Sweetly scented flowers.
J. sp. 680 Herb. Mkonjane in Acacia nigrescens Woodland.
4464 Suregada zanzibarensis Baill. 1117 Tree. Ulukondo. Sand Forest.
4478a Spirostachys africana Sond. 1756 EJM Tree. Mahemane. Frequent in pan-
edge communities; on boulder beds. Antelope and monkeys eat dried fallen
leaves. Stingless bees use these trees extensively for their nests.
4483 Sapium ellipticum (Hochst.) Pax 1342 Tree. Pongola Riverine Forest.
S. integerrimum (Hochst. ex Krauss) J. Leon 312 Tree. Ulukondo. Sand
Forest.
4498 Euphorbia cooperi N.E.Br. ex Berger var. cooperi * Succulent shrub. Aber-
corn drift. Planted in camp and crocodile farm gardens.
E. geniculata Orteg. 404 Herb. Usutu Riverine Forest.
E. grandicornis Goebel ex N.E.Br. subsp. grandicornis 646 Succulent shrub.
Mahemane. Thicket, disturbed areas, particularly round old kraal sites between
Banzi and Inyamiti pans. Attracts many insects when in flower, especially wasps.
Monkeys eat fruits. Thorns raise painful, itchy bump on skin.
E. hirta L. 559KLT ‘‘Common weed on floodplain’’. (KLT).
E. hypericifolia L. 145 Herb. Pongola Floodplain margins. _
E. Fe eae ex Boiss. Sareulent tree. Mahemane clearing. Widespread.
Monkeys eat flowers and fruits.
E. knuthii Pax 291 Succulent. Mahemane. Common, widespread. i
E. pseudotuberosa Pax 121 Bulb. E slope Ndumu Hill. Uncommon. In Acacia
nigrescens Thicket.
E. tirucalli L. 66 Succulent s
Monkeys eat fruits.
hrub, Ulukondo Sand Forest. Black rhino browse.
36
Journal of South African Botany
Euphorbia triangularis Desf. * Succulent shrub. Abercorn drift, in Lebombo
foothills just outside W boundary. Planted in gardens.
4500 Synadenium cupulare (Boiss.) L. C. Wheeler 1048 KLT Shrub. Ulukondo.
Sand Forest and thicket. Sap causes severe irritation, raising blisters on skin and
causing headache and nausea.
4503 Monadenium lugardae N.E.Br. 67 Succulent. Mahemane—infrequent fairly
large communities.
(107) ANACARDIACEAE
4545 Mangifera indica L. Tree. Frequent, very large specimens in Riverine Forests—
sometimes large communities. Eaten by man and monkeys (introduced).
4558 Sclerocarya caffra Sond.’ 273 Tree. Ndumu Hill. Common locally in wood-
land; occasionally scattered in woodland elsewhere. Spring leaves appear with the
flowers. Flowers cup-shaped when first open, petals fold back later, do not close
at night. Many insects, especially bees, attracted to flowers. Fruits drop when
green, ripen on ground. Eaten by animals and man (mGanu). A beer is brewed
from the fruits by the Tongas. Nuts from the hard seed are used as food by the
Tongas (uMongo, small but delicious, like a walnut). Bark of the trees is used to
dye fish nets. Leaves browsed by antelope. Large caterpillars, found in quantities
on S. caffra in mid-summer, are collected and eaten by the Tongas. A large borer
grub is also collected to eat, by tapping the bark of the tree until a hollow sound
indicates a borer’s hole. Trees can be killed by infestations of this large insect.
When in green fruit, some trees become infested by caterpillars which devour all
the leaves off the tree, and cover the entire tree, trunk and branches, with fine,
strong, silky web. This does not apparently adversely affect the tree, which
produces new leaves and soon returns to normal again. It would be useful to have
the correct names of the caterpillars, borers and moths associated with this tree.
4563 Lannea stuhlmannii (Engl.) Engl. 294 Tree. Ndumu Hill. Woodland. Fruits
eaten by monkeys and Tongas (umGanunkomo). Bark of trees used to dye fish
nets.
4589 Ozoroa obovata (Oliv.) R. & A. Fernandes var. obovata 378 Tree. Ndumu
Hill. Woodland. Browsed.
4594 Rhus fraseri Schonl. 1230 Shrub. Usutu Floodplain.
R. guenzii Sond. var. spinescens (Diels) R. & A. Fernandes 835 Shrub.
Widespread. Browsed by nyala. Birds feed on fruits which are also eaten by
Tongas (umPhondo).
R. sp. 1044 Shrub. E slope Ndumu Hill. Frequent.
(109) CELASTRACEAE (Incl. Hippocrataceae)
4626 Maytenus heterophylla (Eck\. & Zeyher) N. Robson 886 Shrub. Widespread.
Flowers sweetly, sometimes unpleasantly, scented. Blooms up to four times a
season. Heavily browsed.
1646—Ndumu Hill—this form common on hill; leaves grey, black striated bark,
main stem clearly formed and tree-like rather than shrub-like. A very variable
species.
M. nemorosa (Eckl. & Zeyher) Marais 1431 Shrub. Mdipine. Usutu Flood-
plain.
M. senegalensis (Lam.) Exell 825 Shrub. Ndumu Hill. Woodland. Browsed.
Monkey eat leaves and shoots.
Checklist of Plants in Ndumu Game Reserve 37
4628 Putterlickia verrucosa (E. Mey. ex Sond.) Szyszyl. 1415 Shrub. E of Pongola
Floodplain.
464 Cassine aethiopica Thunb. 675 Tree. Ndumu Hill, scattered in woodland.
Browsed. Fruits eaten by Tongas (umGunguluzampunzi).
C. transvaalensis (B. Davy) Codd 799 Tree. Mahemane. Browsed.
4661 Hippocratea africana (Willd.) Loes. var. richardiana (Cambess.) N. Robson
881 Climber. Mahemane. Thicket.
4662 Salacia kraussii (Harv.) Harv. 889 Shrub. Ndumu Hill, woodland. Very com-
mon. Browsed. Fruits eaten by animals and man (amaBonsi). Not affected by fire.
S. leptoclada Tul. 756 Shrub. Ulukondo. Sand Forest. Fruits eaten by Tongas
(uHlangahomo). Browsed.
(110) ICACINACEAE
4709 Pyrenacantha kaurabassana Baill. 128 Climber, with large swollen under-
ground root. Woodland. Fruits eaten by Tongas (inZema).
(111) SAPINDACEAE
4726 Cardiospermum halicababum L. 402 Climber. Common on floodplains.
4734 Allophylus decipiens (Sond.) Radlk. 1417 Shrub. Riverine Forests. Scented
flowers.
4746 Deinbollia oblongifolia (E. Mey.) Radlk. 594 Shrub. Widespread, not com-
mon. Browsed.
4769 Haplocoelum gallense (Engl.) Radlk. 735 Tree. Ulukondo, Sand Forest.
4784 Pappea capensis Eckl. & Zeyher 1435 Tree. Mahemane. Browsed. Fruits
eaten by animals and man (umQuokwane).
(115) RHAMNACEAE
4861 Ziziphus mucronata Willd. 383 Tree. Ndumu Hill. Woodland. Browsed. Bark,
leaves and fruits are eaten by many animals. Tongas eat the berries (umPhafa).
4868 Berchemia discolor (Klotzsch) Hemsl. 416 KLT E of Pongola Floodplain.
Monkeys eat the fruits. :
B. zeyheri (Sond.) Grubov 388 Tree. Ndumu Hill. Woodland. Many insects
visit flowers, especially bees. Heavily browsed. Fruits eaten by birds, monkeys,
antelope and people (umNeyi).
(116) HETEROPYXIDACEAE 1s.
4908a Heteropyxis natalensis Harv. 873 KLT Paphekulu stream. Drainage line in
west.
(117) VITACEAE
4917 Rhoicissus digitata (L.f.) Gilg & Brandt 922 Climber. Ndumu Hill. Wood-
land. Browsed.
R. tridentata (L.f.) Wild & Drummond 1274 Climber. Mahemane Thicket.
Browsed.
4918 Cissus quadrangularis L. 81 Succulent climber. Ndumu Hill, Mahemane.
Common in thicket. ; ; 2
C. rotundifolia (Forsk.) Vahl 114. Succulent climber. Ndumu Hill. Frequent in
woodland. Browsed by nyala.
38 Journal of South African Botany
4918a Cyphostemma barbosae Wild & Drummond 677 Succulent herb with large
swollen root. Mkonjane. Not common.
C. cirrhosum (Thunb.) Desc. ex Wild & Drummond subsp. transvaalense (Szy-
szyl.) Wild & Drummond 1240 Succulent climber. Mahemane Thicket.
C. subciliatum (Bak.) Desc. ex Wild & Drummond 451 Climber. Riverine
Forest and Floodplain margins.
(118) TILIACEAE
4953 Corchorus asplenifolius Burch. 1272 Herb. Mahemane Thicket.
C. junodii (Schinz) N.E.Br. 894 Herb. Ndumu Hill.
4966 Grewia bicolor Juss. 807 Shrub. Mahemane. Widespread. Browsed. Fruits ‘|
eaten by birds and Tongas (uSipane).
G. caffra Meisn. 744 Shrub. Widespread. Browsed. Birds, monkeys and |
people eat the fruits (iPhata). Angled stems used for making traps (for monkeys |
and crocodiles etc.).
G. flava DC. 1066a Shrub. Ndumu Hill. Woodland.
G. microthyrsa K. Schum. ex Burret 734 Shrub. Ulukondo. Sand Forest.
Browsed. Birds, monkeys, Tongas eat fruits (uMunyuane).
G. monticola Sond. 330 Tree. Ndumu Hill. Woodland.
G. occidentalis L. 274 Shrub. Mahemane near Banzi pan. Browsed.
G. subspathulata N.E. Br. 410 KLT.
G. villosa Willd. 668 Shrub. Mkonjane. Acacia nigrescens Woodland.
4975 Triumfetta pentandra A. Rich. 507 Herb. Ndumu Hill. Woodland.
(119) MALVACEAE
4983 Abutilon austro-africanum Hochr. 83a Herb. Pongola Floodplain margin.
A. engleranum Ulbr. 148 Herb. Ndumu Hill. Woodland.
A. grandiflorum G. Don 1359 Herb. Floodplains. |
A. guineense (Schum.) Bak.f. & Exell 670 Herb. Mkonjane. Acacia nigres- |
cens Woodland. Flowers remain open all day. |
4995 Malvastrum coromandelianum (L.) Garcke 758 Herb. Usutu Floodplain.
4998 Sida cordifolia L. 745 Herb. Ndumu Hill. Woodland. Flowers open early
morning, close midday.
S. dregei Burtt Davy 144 Herb. Pongola Floodplain. Flowers close at midday. |
S. rhombifolia L. 399 Shrub. Floodplains.
5007 Pavonia leptocalyx (Sond.) Ulbr. 751 Herb. Ulukondo. Sand Forest. |
P. patens (Andr.) Chiov. 751 Herb. Mahemane. Thicket. |
5013 Hibiscus calyphyllus Cay. 1314 Shrub. Floodplain margins. Widespread. Well \
browsed.
H. cannabinus L. 472 Herb. Pongola Floodplain. Flowers close midday.
H. dongolensis Del. 539 Shrub. Mkonjane.
H. micranthus L.f. 110 Herb. Pongola Floodplain margins. Flowers open
white, turn red later.
H. palmatus Forsk. 354 Herb. Near Banzi pan, in thicket.
H. pusillus Thunb. 1006 Herb. Ndumu Hill. Woodland; fairly widespread.
Flowers can be yellow or mauve and usually open after rain or storm; open early
morning, close in afternoon.
H. sabiensis Exell 110a Herb. Ulukondo. Sand Forest.
H. schinzii Guerke 746 Herb. Ndumu Hill. Observed in and near the main
camp.
5018
5019
5020
(120)
5047
5053
5056
5059
5083
5091
(121)
Si
(122)
5199
(125)
5271
(126) FLACOURTIACEAE
5284 Oncoba spinosa Forsk.
Checklist of Plants in Ndumu Game Reserve 39
Hibiscus surattensis L. 458 Scandent shrub. Usutu Floodplain. Flowers close
in afternoon.
H. trionum L. 1462 Herb. Pongola Floodplain margins.
H. vitifolius L. subsp. vulgaris Brenan & Exell 471 Shrub. Pongola Floodplain
margins. Flowers close in evening.
Thespesia acutiloba (Bak.f.) Exell & Mendonca 31 Tree. Ndumu Hill; Pon-
gola Riverine Forest. Flowers close mid-afternoon. Fruits eaten by birds. Browsed
by antelope.
Cienfuegosia hildebrandtii Garcke 1390 Shrub. Mkonjane. Acacia nigrescens
Woodland. Heavily browsed.
Gossypium herbaceum L. var. africanum (Watt) Hutch. & Ghose 1488 Scan-
dent shrub. Pongola Floodplain margins.
STERCULIACEAE
Melhania didyma Eckl. & Zeyher 1033 Herb. Ndumu Hill. Woodland. Flo-
wers open in afternoon, close at night. Scented.
M. forbesii Planch. ex Mast. 1296 Herb. Common, west of Shokwe. Wood-
land.
Dombeya rotundifolia (Hochst.) Planch. var. rotundifolia 830 Tree. Usutu
Floodplain. Uncommon. Sweetly scented flowers.
Hermannia micropetala Hary. 498 Herb. Ndumu Hill. Woodland.
Waltheria indica L. 1407 Shrub. Pongola Floodplain margins.
Sterculia rogersii N.E.Br. 23 Tree. Common on Ndumu Hill. Woodland.
Fruits resemble a peach, with the velvety pink/grey/green bloom. Seeds eaten by
birds and Tongas, but hairs in the pod cause itchy irritation to skin (nKhumphenk-
huphe). Transplants easily from truncheons. Browsed.
Cola microcarpa Brenan 867 Tree. Ulukondo. Sand Forest.
OCHNACEAE
Ochna arborea Burch. ex DC. 697. Shrub. Ndumu Hill. Woodland. No leaves
with flowers (scented). Fruits orange rather than red; eaten by monkeys.
O. natalitia (Meisn.) Walp. 135b Shrub. Ndumu Hill. Woodland. Flowers
appear with leaves, scented. New leaves bright coppery red. Very pretty bright
deep-pink galls on this plant are conspicuous. Browsed.
O. serrulata (Hochst.) Walp. 14 ICP
CLUSIACEAE (nom. alt. Guttiferae) :
Garcinia livingstonei T.And 711 Tree. Ndumu Hill. Woodland. Fruits appear
on old wood, very delicious, eaten by animals and man (umPhimbu). Browsed.
VIOLACEAE
Hybanthus enneaspermus (L.) F. Muell. var. serratus Engl. 111
gola Floodplain.
Herb. Pon-
1065 Tree. Riverine Forests, occasional on drainage
lines.
40 Journal of South African Botany
5284a Xylotheca kraussiana Hochst. var. kraussiana 422 Shrub. Ndumu Hill. In
thickets in woodland. Flowers scented. Petals drop after four days. Fruits split
into segments to expose red and black sticky seeds, eaten by birds, Tongas.
Browsed.
5304 Scolopia zeyheri (Nees) Szyszyl. 1422 Shrub. Pongola Floodplain margins.
5328 Dovyalis caffra (Hook.f. & Harv.) Hook.f. 763 Tree. Mahemane. Widespread.
Many insects attracted to flowers. Fruits eaten by animals and man. Browsed
(umQokolo).
D. longispina (Harv.) Warb. 286 Shrub. Near Shokwe pan, in thicket. Birds,
Tongas eat the fruits (umNyazuma).
(128) PASSIFLORACEAE
5370 Adenia hastata (Harv.) Schinz 1156 Climber. Mahemane Thicket.
A. senensis (K1.) Engl. 862 Climber. Ulukondo. Sand Forest.
(131) CACTACEAE
Opuntia ficus-indica L. Mill. 78 TBO Succulent weed. NRC house surrounds
and scattered in the reserve (introduced).
O. megacantha NRC house and scattered elsewhere in the reserve, despite
attempts to eradicate both species (introduced).
(133) THYMELACACEAE
5435a Lasiosiphon capitatus (L.f.) Burtt Davy 667 Shrub. S boundary in the west,
Acacia nigrescens Woodland.
5442 Synaptolepis kirkii Oliv. 109 Climber. Ndumu Hill. Forms occasional thickets.
Scented.
(134) LYTHRACEAE
5480 Galpinia transvaalica N.E.Br. 366 Tree. On slopes of Ndumu Hill. Scented
flowers attractive to insects. Flowers only last a few days but make the tree
conspicuous for that brief time.
(137) COMBRETACEAE
5538 Combretum acutifolium Exell 5362 EJM_ Tree. Ulukondo. Locally common.
Sand Forest.
C. apiculatum Sond. subsp. apiculatum 786 Tree. Ulukondo. Sand Forest.
C. collinum Fresen. subsp. suluense (Engl. & Diels) Okafor 1116 Tree.
Ulukondo; Ndumu Hill. Sand Forest, woodland. Flowers attract many insects.
C. hereoense Schinz 69 Tree. Common in Acacia nigrescens Woodland.
Browsed.
C. imberbe Wawra 1066 Tree. Scattered in Acacia nigrescens Woodland in
west and on margins of Ndumu Hill.
C. kraussii Hochst. 1665 Tree. Ulukondo. Sand Forest.
C. microphyllum Klotzsch. 828 Climber. Floodplains. Browsed. Climber to
tops of trees where it flowers and fruits prolifically.
C. molle R.Br. ex Don 919 Tree. Ndumu Hill. Woodland. Browsed.
C. zeyheri Sond. 676 Tree. Ndumu Hill. Woodland.
5539 Pteleopsis myrtifolia (Laws.) Engl. & Diels 789 Tree. Ulukondo. Browsed.
Common in Sand Forest.
Checklist of Plants in Ndumu Game Reserve 41
5541 Quisqualis parviflora Gerrard ex Harv. 1100 Climber. Ulukondo. Common in
Sand Forest.
5544 Terminalia phanerophlebia Engl. & Diels 100 TPD Paphekulu stream.
Drainage line in SW corner of reserve.
T. sericea Burch. 419 Tree. Common, Ndumu Hill. Woodland. Unpleasantly
scented flowers, attracts many insects. Browsed. Very hard, ant-resistant wood;
makes excellent fire-wood.
(138) MYRTACEAE
5578 Eugenia mossambicensis Amshoff 803 Shrub. Common on Ndumu Hill.
Woodland. Forms thickets. Not affected by fire. Browsed. Flowers scented, attract
many insects. Fruits eaten by animals and Tongas (iNontsane).
5583 Syzygium guineense (Willd.) DC. 2029 CJW_ Tree. Floodplains. Browsed.
Monkeys eat flower buds, flowers, fruit. Tongas eat fruits (umDoni-wamanzi).
Psidium guajava L. Tree. Large groves along river banks (introduced).
(140) ONAGRACEAE
5793 Ludwiggia octovalvis (Jacq.) Raven subsp. octovalvis 493 Herb. Floodplains.
At Banzi pan it grows in the raft of floating vegetation. Flowers open and close in
the morning.
L.. stolonifera (Guill. & Perr.) Raven 659 Semi-emergent aquatic; Pongola
Floodplain. On pans and seasonal pools. Flowers close in early afternoon.
(141) TRAPACEAE
5829 Trapa natans L. var. bispinosa (Roxb.) Makino 320 Aquatic. Common in
Banzi pan. Spiny fruits collected, taken home and dried, then cracked open and
eaten by Tongas (iniYva). Food for waterfowl too.
(144) ARALIACEAE
5872 Cussonia arenicola Strey 65 Shrub. Ulukondo. Sand Forest.
(145) APIACEAE (nom. alt. Umbelliferae)
5894 Centella asiatica (L.) Urban 459 Herb. Floodplains.
(150) PLUMBAGINACEAE 1
6343 Plumbago zeylanica L. 599 Shrub. Usutu Floodplain. Infrequent. Faintly
scented. Flowers close at night, re-open next day.
(151) SAPOTACEAE
6368 Sideroxylon inerme L. 879 Tree. Margins of floodplains; Mahemane; drainage
lines. Fruits eaten by birds, monkeys.
6386a Manilkara discolor (Sond.) J. H. Hemsl. 1831 Tree. Ulukondo. Sand Forest.
Tongas eat fruits (umNweba). : ;
Wisieochisia (Bak.) Dubard. 1054 Tree. Ndumu Hill; Mahemane. Common in
thicket. Flowers without leaves, heavy honey scent which attracts many insects.
Browsed. Birds, monkeys, Tongas eat fruits (umNqambo).
(152) EBENACEAE ale ‘
6404 Euclea crispa (Thunb.) Guerke var. crispa 336 Shrub. Ndumu Hill. Commo
shrub in woodland. Monkeys, Tongas eat fruits.
aM bi Hiern 844 Tree. Mahemane. Common. Scented flowers.
Browsed. Fruits eaten by birds, monkeys and Tongas (umHlangula).
42
6406
(153)
6440
(154)
6444
6446
(155)
6460
Journal of South African Botany
Euclea natalensis A.DC. 1404 Tree. E of Pongola Floodplain.
E. schimperi (A.DC.) Dandy var. daphnoides (Hiern) de Winter 767 Tree.
Mahemane Thicket. Common. Browsed.
E. undulata Thunb. var. myrtina (Burch.) Hiern 1129 Tree. Mahemane.
Scented, many insects. Fruits eaten by monkeys.
E. undulata Thunb. var. undulata 766 Tree. Mahemane Thicket. Scented.
Monkeys eat fruits.
Diospyros galpinii (Hiern) de Winter 768 Herb. Ndumu Hill. Woodland.
D. lycioides Desf. subsp. guerkei (Kuntze) de Winter 651 Shrub. Ndumu Hill.
Woodland. Forms small thickets. Citrus-scented flowers.
D. villosa (L.) de Winter 1186 Climber? Mahemane. Common in thicket.
OLEACEAE
Jasminum breviflorum Harv. ex C.H.Wr. 1025 Climber. Ndumu Hill. Wood-
land. Very strong, rather unpleasant scent.
J. fluminense Vell. 1192 Climber. Pongola Floodplain margins. Common.
Flowers heavily scented, close at night to open again next day. Browsed.
J. stenolobum Rolfe 124 Scandent shrub. Ndumu Hill. Woodland. Scented.
Browsed.
J. streptopus E. Mey. var. transvaalensis (S. Moore) Verdoorn 774 Climber.
Ulukondo. Sand Forest.
SALVADORACEAE
Azima tetracantha Lam. 874 Shrub. Widespread in thicket, common.
Browsed.
Salvadora angustifolia Turrill var. australis (Schweick.) Verdoorn 837 Tree.
Mahemane Thicket. Foul-smelling flowers.
LOGANIACEAE
Strychnos decussata (Pappe) Gilg 752 Tree. Mahemane Thicket. Browsed.
S. madagascariensis Poir. 713 Tree. Ndumu Hill. Woodland. Browsed. Forms
occasional low thickets. Monkeys eat fruits. Tongas collect fruits, grind up, dry,
then add sugar and eat (nKwakwa).
S. spinosa Lam. 793 Tree. Ndumu Hill. Woodland. Common. Browsed.
Scented flowers. Fruits eaten by animals and man (Mahlala). In dry seasons
antelope congregate around these trees, scarcely moving, whilst feeding off the
fruits.
1122, 792: Ulukondo plants different—warty fruits, large leathery leaves.
1159: Bunguzane plants also look quite different.
S. usambarensis Gilg 1681 Tree. Ulukondo. Sand Forest.
6469 Nuxia oppositifolia (Hochst.) Benth. 803 KLT Usutu Riverine Forest.
(156)
6484
GENTIANACEAE
Enicostemma hyssopifolium (Willd.) Verdoorn 1343 Herb. Usutu Floodplain.
6545 Nymphoides sp. c.f. N. rautenannii N.E.Br. 1578 Aquatic. Mahemane pools.
(157)
Uncommon, found in shallow water of seasonal pools.
APOCYNACEAE
6558 Acokanthera oppositifolia (Lam.) L. E. Codd 1151 Shrub. Mahemane Thicket.
6559
Carissa bispinosa (L.) Desf. ex Brenan var. acuminata (E. Mey.)
Codd 70 Shrub. Mahemane Thicket. Heavily scented flowers. Browsed. Fruits
eaten by birds, monkeys and Tongas.
Checklist of Plants in Ndumu Game Reserve 43
Carissa tetramera (Sacleux) Stapf 518 Shrub. Mahemane Thicket. Very
strongly scented—fills the air on hot humid summer mornings. Browsed. Fruits
eaten by birds, monkeys, Tongas (uQondo).
6562 Landolphia kirkii Dyer 797 Climber. Ndumu Hill. Very common in thicket in
woodland. Eaten by animals and Tongas (amaBungwa).
L. petersiana (Kl.) Dyer 253 Climber. Ndumu Hill. Woodland. Heavily
scented flowers fill the air for the few days of full flowering. Cover entire tree.
peters drop after a day. Spectacular sight and scent. Monkeys, Tongas eat the
Tuits.
6603 Tabernaemontana elegans Stapf 1193 Tree. Floodplains. Sweetly scented
flowers. Browsed. Monkeys eat leaves, flower buds, flowers, fruit. Tongas eat
fruit (umKashlu). White sap used as a bird lime by young boys when trapping
birds.
6619 Rauvolfia caffra Sond. 876 Tree. Riverine Forests. Scented flowers. Browsed.
Monkeys eat leaves, flower buds, flowers, fruit. Tongas grind up bark and leaves,
soak, then drink liquid to alleviate ‘fever’ (umKhahluvungu).
6680 Adenium obesum (Forsk.) Roem. & Schult. var. multiflorum (Klotzsch) Codd
637 Succulent. Widespread. Large specimens confined to thicket now, due to
pressure of browsing animals.
6681 Pachypodium saundersii N.E.Br. * Succulent. Abercorn drift road, in foothills
of Lebombo west of reserve. Planted in gardens.
6688 Strophanthus gerrardii Stapf 64 Climber. Ulukondo; between Inyamiti and
Banzi pans. Sand Forest and thicket. Stem thickens to produce corky-woody
triangular shaped bark. e)
S. luteolus L. E. Codd 27 Climber. Ulukondo; S$ bank Inyamiti. Sand Forest,
thicket. Flowers open for days before dropping, scented.
6689 Wrightia natalensis Stapf 71 Tree. Ulukondo; Mahemane Thicket. Sand
Forest. Scented flowers. Browsed. Used for spear shafts.
(158) PERIPLOCACEAE
6730 Tacazzea apiculata Oliv. 547 KLT Pongola Riverine Forest.
6740 Cryptolepis obtusa N.E.Br. 489 Climber. Usutu Riverine Forest.
6741 Stomatostemma monteiroae (Oliv.) N.E.Br. 614 Climber. Mahemane. Very
common in thicket.
6747 Raphionacme flanaganii Schltr. 885 Climber. Mahemane Thicket.
(159) ASCLEPIADACEAE hae
6778 Schizoglossum garcianum Schltr. ex descr. 122 Herb. Ndumu Hill, woodland.
6791 Asclepias fruticosa L. 1275 Herb. Mahemane Thicket.
A. stash (E. Mey.) Schltr. 1814 Herb. Pongola Floodplain. Has prolifer-
ated in recent years to become very common, forming fields.
6810 Pentarrhinum insipidum E. Mey. 461 Climber. Uncommon on Ndumu Hill.
6834 Cynanchum mossambicense K. Schum. 533 Climber. Uncommon on Ndumu
Hill. Woodland.
44 Journal of South African Botany
6849 Sarcostemma viminale (L.) R.Br. 55 Succulent climber. Common and wide-
spread. Flowers have attractive scent. Browsed by animals. Tongas eat green fruits
and new shoots (umPelepele).
6860 Secamone delagoensis Schltr. 791 Climber. Ulukondo. Sand Forest. Flowers
heavily scented.
S. parvifolia (Oliv.) Bullock 317 Climber. Ndumu Hill. Woodland. Very
strong honey-like scent. Browsed.
6874 Ceropegia monteiroae Hook. 510 Succulent climber. Ulukondo. Sand Forest
(1st record for Natal).
C. plicata E. A. Bruce 460 Succulent climber. Ndumu Hill, widespread but
not common. Flowers drop after a couple of days. ‘
C. rendallii N.E.Br. 422 Succulent climber. Ndumu Hill. Woodland.
C. sandersonii Hook.f. 510 Succulent climber. Ulukondo. Sand Forest. Flo-
wers are scented, droop after a couple of days. Stingless bees feed off flowers.
C. stapeliiformis Haw. var. serpentina (Bruce) H. Huber 423 Succulent climb-
er. Ulukondo. Sand Forest.
C. stapeliiformis Haw. var.? 424 Succulent climber. Ulukondo. Sand Forest.
6875 Riocreuxia torulosa (E. Mey.) Decne. 456 Climber. Usutu Floodplain. Pro-
fusion of this plant in reedbeds on edge of forest. Heavy wild-honey scent. |
6885 Stapelia gigantea N.E.Br. 8 JVH Succulent. Mahemane. Widespread.
Browsed by antelope.
S. longidens N.E.Br. 505 Succulent. Ndumu Hill. Woodland. |
6887 Huernia hystrix N.E.Br. 101 Succulent. Mahemane Thicket.
H. zebrina N.E.Br. 283 Succulent. Ndumu Hill. Woodland.
6891 Gymnema sylvestre (Retz.) Schultes 457 Climber. Usutu Floodplain. Flowers |
scented, attract many butterflies. j
6899 Tylophora lycioides Decne. 1209 Climber. Usutu Floodplain. :
6911 Marsdenia floribunda (E. Mey.) N.E.Br. 77 Climber. Mahemane Thicket.
6917 Pergularia daemia (Forsk.) Chiov. 496 Climber. Common on Ndumu Hill.
Woodland. Flowers open in late afternoon.
(160) CONVOLVULACEAE
6973 Evolvulus alsinoides (L.) L. var. linifolius (L.) Bak. 1308 Herb. Mahemane
Thicket.
E. nummularis L. 761 Herb. Banzi pan shoreline. Small prostrate plant that
creates a carpet of green over the dry black clay.
6978 Seddera capensis (E. Mey. ex Choisy) Hall. 678 Herb. S boundary in west.
S. suffruticosa (Schinz) Hall.f. 430 Herb. Mahemane Thicket.
6991 Jacquemontia tamnifolia (L.) Griseb. 517 Climber. Pongola Floodplain mar-
gins. Flowers close mid-morning.
6993 Convolvulus farinosus L. 640 Climber. Floodplains.
6995 Hewittea sublobata (L.f.) Kuntze 1282 Climber. River-banks.
6997 Merremia palmata Hall.f. 88 Climber. Ndumu Hill. Woodland.
M. tridentata (L.) Hall.f. subsp. angustifolia (Jacq.) Ooststr. 1560 Climber.
Mahemane Thicket.
7003
(161)
7038
7043
7052
7056
Checklist of Plants in Ndumu Game Reserve 45
Ipomoea albivenia (Lindl.) Sweet 40 Climber. Ndumu Hill. Flowers open at
night, close at midday—scented. Flowers and leaves attacked by caterpillars.
Large underground rootstock. Seeds wrapped in ‘cotton wool’; when pods open
to reveal them, creeper appears to be in flower again. Blooms prolifically for 3 to
4 weeks. Leafless in winter.
I. bolusiana Schinz 1666 Climber. Ndumu Hill. Woodland.
I. cairica (L.) Sweet 770 Climber. Ulukondo; Pongola river banks; Usutu
Riverine Forest. Very common. Flowers close mid-afternoon. Monkeys eat
shoots, flower buds, flowers, fruits. Hippo observed eating bunches of the creeper.
I. coptica (L.) Roth ex Roem. & Schult. 1562 Climber. Seepage lines on
Ndumu Hill.
I. digitata L. var. digitata 324 Climber. Pongola Riverine Forest. Flowers up
to height of 20 m, rooted in deep shade, large rootstock.
I. digitata L. var. eriosperma (P. Beauv.) Rendle 323 Climber. Pongola
Riverine Forest. Flowers up to height of 20 m. Woody main stem. Large
rootstock. Flowers open early morning, close early afternoon.
I. lapathifolia Hall.f. 1567 Climber. Balemhlanga. Drainage line in west (1st
record for Natal).
I. magnusiana Schinz 776 Climber. Ndumu Hill. Flowers close at midday.
I. obscura (L.) Ker-Gawl. var. fragilis (Choisy) A. Meeuse 1233 Climber.
Ndumu Hill. Woodland.
I. pellita Hall.f. 143 Climber. E slope Ndumu Hill.
I. sinensis (Desr.) Choisy subsp. blepharosepala (Hochst. ex A. Rich.) Verdc.
1617 Climber. S boundary in west.
I. wightii (Wall.) Choisy 490 Climber. Usutu Floodplain. Flowers open early,
close midday.
BORAGINACEAE
Cordia caffra Sond. 1246 Tree. Usutu Floodplain. Fruits eaten by Tongas
(iLovu amakhulu). : ;
C. ovalis R.Br. 814 Shrub. Mahemane, common and widespread. Fruits eaten
by Tongas (iLovu mcani).
Ehretia amoena Klotzsch 301 Tree. Pongola Floodplain margins. Flowers
sweetly scented, attracting many insects. Browsed. Birds, monkeys, Tongas eat
fruits (umHlele amakhulu). ;
E. rigida (Thunb.) Druce 369 Shrub. Ndumu Hill; Mahemane. Widespread.
Flowers sometimes scented. Leaves very variable in size and shape. Much
browsed. Fruits eaten by birds, monkeys, Tongas (umHlele). Used for making
Fonya (thrust) fishing baskets.
Heliotropium ciliatum Kaplan 138b Herb. Ndumu Hill. Woodland.
H. curassavicum L. 822 Herb. Inyamiti pan. =! sae
H. indicum L. 829 Herb. Floodplains. Eaten as ‘spinach (imifino ubhoyo)
by Tongas.
H. ovalifolium Forsk. 581 Herb. Usutu Floodplain.
H. steudneri Vatke 665 Herb. Mahemane Thicket.
Trichodesma zeylanicum (Burm.f.) R.Br. 524 Herb. Usutu Floodplain. S
boundary fence near Agate hill.
(162) VERBENACEAE
7138
Verbena bonariensis L. 403 Shrub. Floodplains (introduced).
46 Journal of South African Botany
7144 Lantana camara L. 227 Scandent shrub. W of Shokwe (introduced).
L. rugosa Thunb. 1491 Shrub. Ndumu Hill. Fruits eaten by birds, monkeys
and Tongas (umKukutwane). Aromatic leaves and stems.
7145a Phyla nodiflora (L.) Greene 425 Herb. Inyamiti causeway at Pongola channel.
Floodplains.
7148 Chascanum hederaceum (Sond.) Moldenke var. natalense (H. H. W. Pearson)
Moldenke 1807 Herb. Ndumu Hill. Woodland.
C. schlechteri (Guerke) Moldenke 205 Herb. Ndumu Hill; Agate hill, south
boundary. Flowers close at night to open again next day.
7153 Priva cordifolia (L.f.) Druce var. abyssinica (Jaub. & Spach) Moldenke
759 Herb. Widespread and common. Leaves cling together. Sticky clinging
fruits.
7185 Premna mooiensis (H. H. W. Pearson) Pieper 1303 Tree. Mahemane; Acacia
tortilis Woodland. Aromatic leaves.
7186 Vitex patula E. A. Bruce 780 Shrub. Common in Ulukondo. Sand Forest.
Browsed. Tongas eat fruits (imBendula).
V. harveyana H. H. W. Pearson 118 Shrub. Paphekulu. Drainage line in west.
Scented. Fruits eaten by birds, monkeys, Tongas (imBendula).
7191 Clerodendron glabrum E. Mey. 1336 Shrub. Floodplains. Very variable
leaves. Browsed. Scented. Flowers attractive to butterflies.
7192 Holmskioldia tettensis (Klotzsch) Vatke 729 Shrub. Lebombo foothills—
planted in gardens.
(163) LAMIACEAE (nom. alt. Labiateae)
7236 Acrotome hispida Benth. 848 Herb. Ulukondo. Sand Forest.
7264 Leonotis nepetifolia (L.) Ait.f. 597 Herb. Usutu Floodplain.
7268 Leucas glabrata (Vahl) R.Br. ex Benth. 50 Herb. Common, widespread,
browsed.
7281 Stachys spathulata Burch. ex Benth. 522 Herb. Usutu Floodplain.
7342 Hyptis pectinata (L.) Poit. 601 Herb. Usutu Floodplain. Common. Also com-
mon in disturbed areas E of Pongola Floodplain.
7350 Plectranthus amboinicus (Lour.) Spreng. 107 Herb. Mahemane. Leaves aro-
matic.
P. cylindraceus Hochst. ex Benth. 521 Succulent herb. S bank Inyamiti pan.
Thicket.
P. neochilus Schltr. 45 Slightly succulent herb. Mahemane. Very common.
Aromatic, sticky to touch.
P. spicatus E. Mey. ex Benth. 626 Slightly succulent herb. Mahemane
Thicket.
P. vagatus (E. A. Bruce) L. E. Codd 483 Mahemane Thicket.
P. verticillatus (L.f.) Druce 519 Succulent herb. Ulukondo. Sand Forest.
7357 Hoslundia opposita Vahl var. decumbens (Benth.) Bak. 154 Herb. Ndumu
Hill. Browsed. Fruits eaten by animals and Tongas, delicious (uYaweyawe).
7357a Iboza riparia (Hochst.) N.E.Br. * Succulent shrub. Lebombo foothills, planted
in gardens.
Checklist of Plants in Ndumu Game Reserve 47
7364 Basilicum polystachyon (L.) Moench 484 Herb. Floodplains.
7366 Ocimum canum Sims 343 Herb. Mahemane Thicket.
O. urticifolium Roth 1298 Herb. Near Shokwe pan.
7366a Becium obovatum (E. Mey. ex Benth.) N.E.Br. 116 Herb. Ndumu Hill.
Woodland.
7367 Orthosiphon suffrutescens (Thonning) J. K. Morton 123 Herb. Ndumu Hill,
woodland.
(164) SOLANACEAE
7377 Nicandra physaloides Gaertn. 592 Herb. Ndumu Hill, Usutu Floodplain.
7379 Lycium acutifolium E. Mey. 1140 Shrub. Mahemane Thicket. Browsed.
L. oxycladum Miers. 3149 CJW_ Shrub. NW margin of Inyamiti.
7400 Withania somnifera (L.) Dunal 1187 Shrub. Usutu Floodplain. On disturbed
ground. ;
7407 Solanum capense L. 142 Climber. Frequent and widespread.
S. coccineum Jacq. 1387 Shrub. Mahemane Thicket.
S. incanum L. 816 Shrub. Inyamiti pan margins.
S. nigrum L. sens. lat. 309 Herb. Ndumu Hill. Woodland.
S. panduraeforme E. Mey. 1334 Shrub. Very common, floodplains, Ndumu
Hill. Leaves, fruits browsed.
(165) SCROPHULARIACEAE
7517 Manulea parviflora Benth. 596 Herb. Usutu Floodplain. Uncommon.
7519 Sutera brunnea Hiern var. macrophylla Hiern 227a Herb. F slope Ndumu
Hill, Acacia nigrescens Thicket.
7564 Ilysanthes dubia (L.) Bernh. 1557 Herb. Mahemane pools, in shallow water.
7597a Alectra kirkii Hemsl. 525 Herb. Usutu Floodplain. Flowers close at night,
re-open in morning.
7611 Buttonia superba Oberm. 1357 Climber. Mahemane Thicket. Flowers open
deep pink, fade to white over the days. Colour very variable.
7614 Graderia scabra Benth. Herb. Agate hill, SW boundary fence.
7625 Striga forbesii Benth. 47 Herb. Common on Pongola Floodplain.
S. gesneroides (Willd.) Vatke ex Engl. 627. Herb. Mahemane Thicket margins
of floodplains.
(167) BIGNONIACEAE ,
7713 Tecomaria capensis (Thunb.) Spach 645 Shrub. Margins of Pongola Flood-
plain. Infrequent.
7722 Rhigozum zambesiacum Bak. 120 Shrub. Widespread in thicket, but not com-
mon. Forms thickets occasionally. Spectacular sight after first rains—profuse
blooming of yellow flowers. Lovely scent. Flowers eaten by beetles. Browsed by
antelope.
iochi 7 in. Flowers eaten
7161 Kigelia africana (Lam.) Benth. 1480 Tree. Usutu Floodplain
off Bae by antelope. Flowers appear with new leaves. Monkeys eat flower
buds and flowers. Browsed.
48
(168)
VT
7778
7780
(170)
7901
(171)
7914
7939
7965
7972
7973
7978
7980
7985
8007
8031
8048
8094
Journal of South African Botany
PEDALIACEAE
Sesamum alatum Thonn. 445 Herb. Ndumu Hill. Common on disturbed
ground.
Ceratotheca triloba (Bernh.) Hook.f. 584 Herb. Usutu Floodplain. Found on
roadsides. On S boundary in west, flowers very much smaller and deeper
pink—ESP 1612.
Dicerocaryum zanguebarium (Lour.) Merrill 1333 Climber. Usutu Flood-
plain. Long, prostrate stems. Nyala eat leaves and fruits.
LENTIBULARIACEAE
Utricularia inflexa Forsk. var. inflexa 590 Aquatic. Banzi pan.
U. inflexa Forsk. var. stellaris (L.) P. Tayl. 1576 Aquatic. Seasonal pools.
U. reflexa Oliver 1686 Aquatic. Channel across road to the bifurcation of the
Pongola river.
ACANTHACEAE
Thunbergia dregeana Nees 1810 Climber. Ndumu Hill. Common in wood-
land.
Dyschoriste depressa Nees 226 Herb. Pongola Floodplain.
Ruellia patula Jacq. 208(Blue) 63(White) Herb. Common and widespread.
Crabbea nana Nees 1386 Herb. Ndumu Hill; Acacia nigrescens Woodland in
west.
Barleria crossandriformis C.B.Cl. 482 Scandent shrub. Mahemane Thicket.
B. delagoensis Oberm. 769 Shrub. Ulukondo. Sand Forest.
B. elegans S. Moore 470 Shrub. Widespread and common especially on
floodplain margins. Seed pods burst open in the heat of the day, propelling the
seeds some distance from the plant.
B. gueinzii Sond. 631 Shrub. Mkonjane, in thicket on rhyolite outcrops.
Schlerochiton caeruleus (Lindau) S. Moore 346 Scandent shrub. Ulukondo.
Common in Sand Forest. Scented flowers. Browsed.
Blepharis integrifolia (L.f.) E. Mey. ex Schinz 1176 Herb. Mahemane
Thicket.
B.sp. 1299
Crossandra fruticulosa Lindau 43 Herb. Mahemane. Common in thicket.
Flowers over a long period. Browsed.
C. greenstockii S. Moore 547 Herb. Agate hill, on S boundary fenceline in
west.
Asystasia gangetica (L.) T. Anders. 381 Herb. Ndumu Hill. Woodland.
Dicliptera clinopodia Nees 503 Shrub. SW area of reserve. Faintly scented.
Browsed.
D. heterostegia Presl ex Nees 492 Shrub. Usutu Riverine Forest.
Ecbolium amplexicaule S. Moore 49 Herb. Mahemane. Common on disturbed
ground.
Justicia sp. aff. J. anagalloides TY. Anders. 1588 Herb. Mkonjane. Acacia
nigrescens Woodland.
Checklist of Plants in Ndumu Game Reserve 49
Justicia capensis Thunb. 583 Shrub. Between Banzi and Inyamiti pans. Ton-
gas use crushed leaves and stems to obscure human smell after setting snares
(iKokela).
J. flava (Vahl) Vahl 11 Herb. Ndumu Hill. Common and fairly widespread.
J. glabra Roxb. 454 Shrub. Usutu Forest. Common, locally.
df erate (Nees) T. Anders. 1250 Herb. Mahemane Thicket. Browsed by
nyala.
J. sp. 194 Herb. Ndumu Hill. Common.
8094a Monechma debile (Forsk.) Nees 1341 Herb. Usutu Floodplain.
(173) RUBIACEAE
8136/6 Ke ‘autia lasiocarpa Klotzsch var. lasiocarpa 536 Herb. Ndumu Hill. Flowers
open in evening, closing next morning, scented.
K. virgata (Willd.) Bremek. var. virgata 899 Herb. Usutu Floodplain.
8136/14 Agathisanthemum bojeri Klotzsch subsp. australe Brem. var. australe
1237 Herb. Ndumu Hill. Woodland.
A. chlorophyllum (Hochst.) Brem. 4314 EJM Herb. Mahemane Thicket.
8136/18 Pentodon pentander (Schum.) Vatke 956 KLT Herb. East of Banzi pan.
8226 Adina microcephala (Del.) Hiern var. galpinii (Oliv.) Hiern 1683 Tree. Flood-
plains; Riverine Forest. Strong, unpleasant scent to flowers. Bark flakes off in
long strips.
8278 Tarenna barbertonensis (Brem.) Brem. 699 Small tree. Usutu Riverine Forest.
8279 Enterospermum littorale Hiern 878 KLT Shrub. Ulukondo. Sand Forest.
8283 Xeromphis obovata (Hochst.) Keay 191 Shrub. Ndumu Hill. Common in
woodland. Sweetly scented flowers. Browsed. Fruits eaten by Tongas (um-
Khwakhwane).
X: rudis (E. Mey. ex Harv.) L. E. Codd 130 Shrub. Bunguzane. Fairly
widespread, in thicket. Fruits eaten by Tongas. Heavily browsed.
8285 Gardenia amoena Sims 347 Shrub. Ulukondo. Sand Forest. Sweet scent quite
different from the other Gardenia species. Fruits eaten by Tongas (iStempe-
swane). Browsed.
G. cornuta Hemsl. 1183 Tree. Mahemane Thicket. Very common. Scent fills
the air in mornings after rain. Monkeys eat buds and fruits. Bushpigs eat fruits.
Antelope browse and eat fruits. ;
G. spatulifolia Stapf & Hutch. 172 Tree. Ndumu Hill. Woodland. Scented.
Usually bloom after rain. Small groups of trees, widely scattered; fruits eaten by
monkeys and antelope. Browsed.
8293 Oxyanthus latifolius Sond. 1411 Tree. E of Pongola Floodplain.
8308 Tricalysia allenii (Stapf) Brenan var. kirkii (Hook.f.) Brenan 140 Shrub.
Ndumu Hill. Heavy sweet scent.
T. junodii (Schinz) Brenan 784 Shrub. Ulukondo. Sand Forest.
8308a Kraussia floribunda Harv. 163 Tree. Pongola Riverine Forest. Slightly scented
flowers, attract many insects mostly flies. Coppice leaves very large and woe
mature leaves. Browsed. Fruits eaten by birds, monkeys, Tongas (amehlenkosa-
sane).
8351 Vangueria cyanescens Robyns 916 Tree. Ndumu Hill. Woodland. Flowers
i i . Fruits eaten by
scented, attract many insects particularly bees and wasps
antelope, monkeys, Tongas—tefreshing on a hot day. Browsed (umViyo).
50
8352
8352a
8352b
Journal of South African Botany
Vangueria chartacea Robyns 1266 Tree. Ulukondo. Sand Forest. Browsed.
Fruits eaten by Tongas (umViyotshane).
Canthium locuples (K. Schum.) Codd 968 KLT Ulukondo. Sand Forest.
C. obovatum Klotzsch 1110 Tree. Ulukondo. Sand Forest.
C. setiflorum Hiern 754 Shrub. Ulukondo. Sand Forest. Scented. Used for
making Fonya (thrust) fishing baskets. Fruits eaten by Tongas—like raisins
(umBhangwe).
C. spinosum (Klotzsch) Kuntze 1153 Shrub. Mahemane Thicket. Fruits eaten
by Tongas (umVuthwamini). Used for making Fonya baskets.
Plectroniella armata (K. Schum.) Robyns 1678 Shrub. S bank Inyamiti pan.
Dinocanthium hystrix Bremek. 1143 Shrub. Mahemane Thicket. Browsed.
Tongas do not use as firewood because it is used as a fence around graves.
8360 Anycylanthus monteiroi Oliv. 158 Shrub. Ndumu Hill. Fruits eaten by tor-
8383
8464
8473
(176)
8562
8564
8568
8569
8590
8591
toises, antelope, monkeys, Tongas (iStlilotshane).
Pavetta catophylla K. Schum. 198 Shrub. Ndumu Hill. Woodland. Browsed.
Fruits eaten by Tongas.
P. edentula Sond. 1033 KLT Tree. Rhyolite outcrops, Mkonjane. Scented
flowers.
P. graciliflora Bremek. 314 Shrub. Banzi pan margins, in thicket.
P. inandensis Bremek. 350 Shrub. Ulukondo. Sand Forest.
P. lanceolata Eckl. 1102 Tree. Ulukondo. Sand Forest. Browsed.
P. schumanniana F. Hoffm. ex K. Schum. 787 Shrub. Ndumu Hill. Forming
thickets in woodland. Heavy but unpleasant scent. Browsed. Fruits eaten by
Tongas.
P. tristis Bremek. 408 Tree. Usutu Riverine Forest.
Richardia brasiliensis Gomez 1691 Herb. Ndumu Hill. Woodland, on dis-
turbed ground.
Borreria scabra (Schumach & Thonn.) K. Schum. 417 Herb. Ndumu Hill.
Common on disturbed ground.
CUCURBITACEAE
Mukia maderaspatana (L.) M. J. Roem. 1345 - Climber. Floodplains.
Zehneria parvifolia (Cogn.) J. H. Ross ined. 605 Climber. Usutu Floodplain.
Z. scabra (L.f.) Sond. 806 Climber. Mahemane Thicket.
Kedrostis foetidissima (Jacqg.) Cogn. subsp. obtusiloba (E. Mey. ex Sond.)
Meeuse 504 Climber. Ndumu Hill. Woodland.
K. hirtella (Naud.) Cogn. 671 Climber. Mkonjane. Acacia nigrescens Wood-
land in west.
Corallocarpus bainesii (Hook.f.) A. Meeuse 342 Climber. Widespread.
Acanthosicyos naudinianus (Sond.) Jeffrey 795 Climber. Ndumu Hill. Wood-
land.
Momordica clematidea Sond. 414 Climber. Usutu Floodplain. Unpleasant
scent.
M. foetida Schum. & Thonn. 406 Climber. Usutu Forest. Unpleasant scent.
M. involucrata E. Mey. 10 Climber. Ndumu Hill. Flowers have unpleasant
scent, close at night, then drop. Birds, Tongas eat the fruits (uHaelane), which
split open when touched at the pointed base.
8599
8608
8610
8628
(177)
8668
8690
8694
(179)
8734
8751
Checklist of Plants in Ndumu Game Reserve 51
Cucumis myriocarpus Naud. 97b Climber. Pongola Floodplain edge.
Trochomeria hookeri Harv. 948 Climber. Ndumu Hill. Woodland.
T. macrocarpa (Sond.) Hook.f. 1001 Climber. Ndumu Hill. Woodland.
Lagenaria mascarena Naud. ex descr. 509 Climber. Usutu Riverine Forest.
Pleasant scent. Climbs to a great height in forest. Large, heavy fruits hang from
old woody stems.
Coccinia rehmannii Cogn. var. littoralis A. Meeuse 344 Climber. Widespread.
Flowers close at midday.
CAMPANULACEAE
Wahlenbergia undulata DC. sens. lat. 25 Herb. Ndumu Hill. Flowers close at
night, re-open next day.
Sphenoclea zeylanica Gaertn. 1609 Herb. Seasonal pools. Stem becomes
enlarged and spongy below water level.
Lobelia filiformis Lam. var. natalensis (A.DC.) Wimm. 1656 Herb. Ndumu
Hill. Seepage on main road near main gate.
COMPOSITAE
Ethulia conyzoides L. 764 Herb. Pongola Floodplain.
Vernonia centaurioides Klatt 477 Herb. Ndumu Hill. Woodland.
V. cinerea (L.) Less. 658 Herb. Floodplains.
V. colorata (Willd.) Drake 551 Shrub. Floodplains. Strongly scented. Many
butterflies and insects attracted to flowers. Charaxes butterflies attracted to the
oozing sap in great numbers. Browsed.
V. fastigiata Oliv. & Hiern 1598 Herb. Mkonjane. Acacia nigrescens Wood-
land.
V. glabra (Steetz) Vatke 5 Herb. Pongola Floodplain. Flowers open deep
blue/purple—faded to paler colours by next day.
V. oligocephala (DC.) Sch. Bip. ex Walp. but approaching V. natalensis Sch.
Bip. 718 Herb. Ndumu Hill. Woodland.
8795 Ageratum conyzoides L. 1424 Herb. Floodplains (introduced).
8818 Mikania cordata (Burm.f.) B. L. Robinson 635 Climber. Usutu Riverine
8865
Forest. Heavily scented flowers, not pleasantly. Many insects attracted to flowers.
Grangea maderaspatana (L.) Poir. 831 Herb. Floodplains.
8900 Aster squamatus (Spreng.) Hieron 1689 Herb. Ndumu Hill, Pongola Flood-
plain (introduced weed).
8925 Nidorella resedifolia DC. 151 Herb. Ndumu Hill. Woodland.
8926
8936
8939
894]
Conyza bonariensis (L.) Cronquist 1034 Herb. Ndumu Hill. Woodland (intro-
duced weed).
Brachylaena huillensis O. Hoffm. 883 Tree. Ndumu Hill. Used for spear
shafts. ae ;
B. ilicifolia (Lam.) Phillips & Schweickerdt 1421 Tree. Mavilo ‘island’ on
Pongola Floodplain.
Blumea mollis (D. Don) Merrill 119 Herb. Pongola Floodplain.
Pluchea dioscoridis (L.) DC. 612 Shrub. Usutu Floodplain. Aromatic scent to
leaves and stems. Browsed.
52
Journal of South African Botany
8953 Epaltes gariepina (DC.) Steetz 74 Herb. Usutu Floodplain.
8955 Sphaeranthus peduncularis DC. 1307 Herb. Mahemane pools.
8992 Gnaphalium calviceps Fernald 6865 H&B Herb. Usutu Floodplain (introduced
weed).
G. luteo-album L. 310 Herb. Pongola Floodplain (introduced weed).
G. pensylvanicum Willd. 656 Herb. Ndumu Hill. Common on roadsides (in-
troduced weed).
G. polycaulon Pers. 132 HF Herb. Pongola Floodplain (introduced weed).
9006 Helichrysum argyrosphaerum DC. 655 Herb. Ndumu Hill. Woodland.
athrixiifolium O. Hoffm. 823 KLT
candolleanum Buek 608 Herb. Usutu Floodplain.
kraussti Sch. Bip. 607 Shrub. Usutu Floodplain. Common on white sands.
oxyphyllum DC. 215 Herb. Ndumu Hill. Woodland.
pilosellum (L.f.) Less. 215a Herb. Ndumu Hill. Woodland.
rugulosum Less. 214 Herb. Ndumu Hill. Woodland.
sp. c.f. H. setosum Harv. 606 Herb. Usutu Floodplain.
po Sy By Be SB
9090 Geigeria burkei Harv. subsp. valida Merxm 373 Herb. Mkonjane. Acacia
nigrescens Woodland in west.
9130 Acanthospermum hispidum DC. 610 Herb. Usutu Floodplain (introduced
weed).
9146 Ambrosia artemisiifolia L. 544 KLT Herb. Pongola Floodplain (introduced
weed).
9166 Eclipta prostrata (L.)L. 410 Herb. Usutu Floodplain. Young plants’ leaves are
very large, stems have no air bladders. Older plants have large leaves at base,
smaller and narrower above, with air bladders in stems (weed).
9204 Melanthera scandens (Schumach. & Thonn.) Roberty subsp. dregei (DC.) Wild
164 Herb. Pongola Riverine Forest.
9207 Spilanthes mauritiana (Persoon) DC. 575 Herb. Floodplains.
9237 Bidens biternata (Lour.) Merrill & Scherff 1621 Herb. Ndumu Hill (intro-
duced weed).
9282 Flaveria bidentis (L.) Kuntze 306 Herb. Ndumu Hill (introduced weed).
9351 Cotula anthemoides L. 1463 Herb. Pongola Floodplain. Uncommon.
C. australis (Spreng.) Hook.f. 1631 Herb. Ndumu Hill (annual weed).
9411 Senecio barbertonicus Klatt 61 Succulent shrub. Common and widespread.
Sweetly scented flowers. Browsed.
S. deltoideus Less. 593 Succulent climber. Usutu Floodplain. Wild-honey-like
scent.
S. fulgens (Hook.f.) Nichols 15 Succulent. Mahemane Thicket. Browsed.
Swollen rhizome eaten by bushpig.
S. macroglossus DC. 1 Succulent climber. Ulukondo. Sand Forest.
S. madagascariensis Poir. 652 Herb. Ndumu Hill. Petals (rays) fold in at
night, open again next day.
S. pleistocephalus S. Moore 843 Succulent climber. Ndumu Hill. Widespread.
Heavy honey-scented flowers. Corky bark on old stems. Dies back after flower-
ing, re-shoots. Browsed.
S. polyanthemoides Sch. Bip. 760 Herb. Shokwe pan margins.
Checklist of Plants in Ndumu Game Reserve 53
Senecio quinquelobus DC. 531 Succulent climber. Mahemane Thicket.
Sweetly scented. Dies back in early summer, re-shoots.
S. transvaalensis Bolus 500 Herb. Mkonjane. Acacia nigrescens Woodland in
west.
S. viminalis Bremek. 325 Succulent climber. Ndumu Hill. Woodland.
Browsed. Blooms more than once in a good season.
9438 Berkheya zeyheri (Sond. & Harv.) Oliv. & Hiern subsp. rehmannii (Thell.)
Roessler 206 Herb. Agate hill. Acacia nigrescens Woodland in west.
9528 Gerbera ambigua (Cass.) Sch. Bip. Herb. Drainage lines on Ndumu Hill; see
ESP no. 440 from Otobotini.
9595 Sonchus asper (L.) Hill 1504 Herb. Pongola Floodplain (introduced weed).
Tongas use with and as pumpkin shoots, ‘imifino’—chopped up, boiled with
curry and peanuts.
S. oleraceus L. 802 Herb. Pongola Floodplain (introduced weed).
9596 Lactuca indica L. 818 Herb. Pongola Floodplain.
REFERENCES
ANDERSON, J. L. and Pootey, E. S., 1977. Some plants identified in the rumina of Nyala
antelope Tragelaphus angazi in Ndumu Game Reserve. Lammergeyer 23: 40-45.
Compton, R. H., 1966. An annotated check list of the Flora of Swaziland. JI S.
Afr. Bot. Suppl. vol. 6.
De Moor, P. P., Poorey, E. S., NeviLLE, G. and BaricutEvy, J., in press. The
vegetation of Ndumu Game Reserve, Natal. A quantitative physiognomic survey.
Ann. Natal Mus.
Dixon, J. E. W., 1966. Notes on the mammals of Ndumu Game Reserve.
Lammergeyer 6: 24-40.
Dutton, T. P., 1972. Some factors affecting the fluctuating waterfowl popula-
tions on Lake Nyamiti. Unpublished Research project, Field Ecology Course,
University of Rhodesia. pp. 1-97.
Dyer, R. A., 1976. Genera of Southern African Flowering plants. 2 vols:
Department of Agricultural Technical Services.
FosBerG, F. R., 1967. A classification of vegetation for general purposes. In: G.
F. Peterken (ed.), Guide to the check sheet for IBP areas. pp. 73-116. Oxford:
Blackwell Scientific publications.
OaTLEy, T. B., 1964. The probing of Aloe flowers by birds. Lammergeyer 8} (Gp
2-8.
Pootey, A. C., 1968. A short note on the diet of the Vervet monkey Cercopith-
ecus aethiops in Zululand. Lammergeyer 9: 29-31.
Ross, J. H., 1972. Flora of Natal. Mem. bot. Surv. S. Afr. 39.
ScHELPE, E. A. C. L. E., 1970. Pteridophyta. Flora Zambesiaca. London: Crown
Agents for Overseas Governments.
ScoTcHER, J. S. B., 1974. A quantitative assessment of the food Risener oi
Hippopotamus amphibius L. in the Ndumu Game Reserve, Tongaland. ; nee :
lished M.Sc. thesis. Pietermaritzburg: Department of Botany, University of Natal.
pp. 1-190.
TinLEY, K. L., 1964a. Fishing methods of the Thonga tribe in North-eastern
Zululand and Southern Mozambique. Lammergeyer 3 (1): 9-39.
, 1964b. Summary of an ecological survey of Ndumu Game Reserve.
Unpublished roneoed report. Natal Parks Board. pp. 1-42.
54 Journal of South African Botany
ACKNOWLEDGEMENTS
I am indebted to Dr O. M. Hilliard for her enthusiastic encouragement over the
years, for assistance and guidance in the task of collecting plants, and for her
comments on this manuscript. I am very grateful to Dr K. D. Gordon-Gray for the
time she has given to the naming of grasses and sedges, for taking great interest in
my work at all times and for checking this manuscript. I wish to thank Miss L.
Smook, Mrs J. Stewart, Mrs D. Fenn and Mr S. Mthalane, also of the Botany
Department of the University of Natal, for their help. Mr B. L. Burtt, Royal
Botanic Gardens Edinburgh, provided inspiration in the field and from abroad—
and assisted greatly in putting names to some difficult specimens. Dr E. J. Moll,
Dept. of Botany, University of Cape Town, kindly commented on parts of the
manuscript and, with Mrs A. A. Mauve and Dr I. Verdoorn of B.R.I. Pretoria; Dr
J. H. Ross of Melbourne, Australia; Mr C. J. Ward of University of Durban-
Westville; have all been most helpful over the years. Mr C. F. Musil of Natal
Herbarium is thanked for his assistance with queries regarding Lemnaceae. I am
grateful to Dr P. P. De Moor, Res. Inst. for Diseases in a Tropical Environment,
for the opportunity of participating in the vegetation survey.
I wish to thank the Natal Parks Board and the following staff for assistance
and facilities provided: Mr. J. L. Anderson, Mr R. S. Crass, Mr R. Porter, Miss L.
Roy, Dr D. R. M. Stewart, Mr J. Vincent. The Research Section of the Natal Parks
Board very kindly made funds available for the publication of the maps. I am
grateful to many of the past Ndumu staff for drawing attention to plants of
interest, particuiarly to Mr and Mrs T. P. Dutton, Mr and Mrs G. O. Jackson, Mr
and Mrs R. Physick, Mr and Mrs G. W. Schutte and to Mr and Mrs A. J.
Tomkinson. Ex-game-guard Sigia Gumede gave valuable assistance in listing the
edible plants.
Greatest thanks go to my husband Tony, whose interest and assistance, at all
times, encouraged me to bring this work to completion. I am also indebted to my
mother, Mrs E. K. Bond, who looked after our two sons whilst the herbarium
work was carried out.
JIS. Afr. Bot. 44 (1): 55-65 (1978)
HABITAT OF THREATENED PROTEACEAE ENDEMIC TO WESTERN
CAPE COASTAL FLATS
A. V. MILEWSKI
(Rare and Endangered Plant Species Research Group, Bolus Herbarium, Univer-
sity of Cape Town)*
ABSTRACT
Serruria ciliata, Diastella buekii and four species of Leucadendron are among members
of the family Proteaceae endemic to the western coastal flats of the south-western Cape.
These six species are associated with types of Coastal Fynbos or Coastal Rhenosterbosveld.
Their habitats are compared using associated plants (Proteaceae and Restionaceae) as an
expression of environmental conditions.
UITTREKSEL
HABITAT VAN BEDREIGDE PROTEACEAE ENDEMIES AAN DIE WESTELIKE
KAAPSE KUSVLAKTES
Serruria ciliata, Diastella buekii en vier Leucadendron spesies is voorbeelde van lede
van die familie Proteaceae wat beperkte verspreidings op die westelike kusvlakte van
suidwes-Kaapland het. Hierdie ses spesies kom op verskillende tipes kusfynbos of
kusrenosterbosveld voor. Hul habitatte word met mekaar vergelyk op grond van geasso-
sieerde plantsoorte (Proteaceae en Restionaceae) as weerspiecling van omgewingstoe-
stande.
INTRODUCTION
The area occupied by natural vegetation on the western coastal flats of the
south-western Cape (a land-unit, defined by Milewski & Esterhuysen, 1977),
corresponding roughly to the western blocks of Acocks’ [1975] Coastal Rhenoster-
bosveld and Coastal Macchia and the Breérivier valley east to Worcester) has
been severely fragmented by man-made environmental changes. Several naturally
tare plant species restricted to this area survive today only as relict populations
and are threatened with extinction (Rare and Endangered Species Research Group,
unpublished; Milewski & Esterhuysen, 1977). Knowledge of the phytogeographi-
cal relationships and habitats (sensu Whittaker et al., 1973) of these localized
habitat-specific endemics is an urgent prerequisite for their conservation.
This paper attempts a comparison of the habitats of several threatened endemic
species in the family Proteaceae by a qualitative floristic method, using a sample
of the plant taxa associated with each in its natural environment. Assumptions
*Present address: School of Environmental and Life Sciences, Murdoch University,
Murdoch, Western Australia.
Accepted for publication 3rd May, 1977.
55
56 Journal of South African Botany
basic to this exploratory approach, which was similar to that of Milewski and
Esterhuysen (1977) and Milewski (1977) were:
(i) that the set of plant species growing at a site is an adequate and easily
documented reflection of local environment in fynbos; the number of species
in a given stand of this vegetation is relatively large (Taylor, in press),
permitting more detailed characterisation of habitats by a floristic method
than might be possible in vegetation dominated by another flora,
(ii) that data on presence and absence of species are suitable and sufficient for
such characterisation, in view of the high species-diversity in fynbos and the
possibility that the relative importances of species have been changed by
man-made disturbance, and
(iii) that two families characteristic of fynbos (Taylor, in press), the Proteaceae
and Restionaceae, constitute a representative sample of the taxa composing
the vegetation and are sufficient for characterisation of habitats for compara-
tive purposes.
Additional considerations leading to the choice of the Proteaceae as a sample
group were that the study species are members of this family and that most genera
are, or have already been, under taxonomic revision. The Restionaceae were in
turn chosen partly because representatives of the family are invariably present in
and thus characterise fynbos, and partly because the apparent habitat specificity of
members of this large group makes them useful indicators of environment (Taylor,
1969, 1972a, in press; Boucher, 1972; Milewski & Esterhuysen, 1977).
The family Proteaceae is represented on the western flats of the south-western
Cape by twelve species of Leucadendron, six species of Leucospermum, ten
species of Serruria, three species of Diastella and six species of Protea. The
following is a brief outline of the phytogeography of these groups in relation to the
overall phytogeography of these flats.
Leucadendron salignum occupies the major part of the western flats and is
widespread throughout the south-western Cape. The remaining 11 Leucadendron
species occurring on the flats are all endemic, extending neither on to adjacent
mountains nor on to the southern coastal flats east of Hangklip. Leucadendron
floridum, L. thymifolium, L. levisanus, L. verticillatum and L. flexuosum, and the
extinct L. spirale, have or had small, continuous areas of distribution. Leucaden-
dron lanigerum, L. corymbosum, L. cinereum, L. stellare and L. foedum have
wider ranges, those of L. stellare, L. corymbosum and L. lanigerum being disjunct.
In addition to these 12 species, Leucadendron chamelaea, L. linifolium, L.
brunioides, L. rubrum and L. spissifolium extend marginally on to the flats from
their main ranges elsewhere (Williams, 1972).
Three species of Leucospermum, viz. L. parile, L. arenarium and L. tomento-
sum, are endemic to small, continuous areas of distribution on the western flats.
Leucospermum hypophyllocarpodendron and L. rodolentum have wider, disjunct
$$$ ga
Habitat of Threatened Proteaceae ii
ranges centred on, but not restricted to, the flats. Leucospermum grandiflorum has
a small range on part of the flats and adjacent foothills. In addition to these six
species, L. conocarpodendron and L. calligerum extend marginally on to the flats
from their main ranges on the mountains (Rourke, 1972).
Serruria burmannii parallels Leucadendron salignum as a widespread inhabi-
tant of most of the western flats. Serruria ciliata, S. trilopha, S. furcellata, S.
linearis and §. roxburghii are all endemic to the flats, having small areas of
distribution; S$. decipiens has a wider range but appears also to be endemic.
Serruria incrassata, S. aitonii, S. glomerata, S. cyanoides and S. brownii occupy
part of the flats and extend on to the adjacent foothills. In addition, S. vallaris and
possibly also S. fucifolia, both mountain species, extend marginally on to the flats
(Rourke, in prep.).
Diastella parilis and D. buekii have small ranges on the flats adjacent to the
foothills. Diastella proteoides, also endemic, has a wider distribution. In addition,
D. divaricata has a range centred on the mountains and extending marginally on to
the flats (Rourke, 1976).
Protea repens is a widespread member of its genus on the western flats, as are
Leucadendron salignum and Serruria burmannii. Protea pulchra and P. acaulos
also occupy the major part of the flats but extend on to the adjacent mountains.
Protea scolymocephala, P. odorata and P. mucronifolia are endemic, P. scolymo-
cephala having a wide range relative to the other two species. In addition, P:
arborea, P. laurifolia, P. cynaroides, P. lepidocarpodendron and P. macroce-
phala, all mountain species, occur marginally on the flats (Rourke, in prep.).
Studies of Restionaceae (Milewski & Esterhuysen, 1977; Milewski, 1977)
represent virtually the only available phytogeographical information dealing speci-
fically with the western flats. Weimarck (1941), working with preliminary distri-
bution data for a number of sample groups over the entire south-western Cape,
treated this area merely as part of his ‘‘South-Western Centre’’, a phytogeographi-
cal land unit comprising both mountains and flats south and west of the Breede
and Berg Rivers. He minimised the phytogeographical distinctiveness of these flats
in comparison with surrounding mountains and the Bredasdorp limestone flats,
stating: ‘‘The species occurring here have, as a rule, large distribution areas and
are also abundant in a large part of the flats’’, and citing Corymbium SALA
Echiostachys spicatus and Lobostemon paniculaeformis as examples of the “‘very
few’’ endemic species (Weimarck, 1941).
The restriction of three Restionaceous (Milewski & Esterhuysen, 1977) and 23
Proteaceous (above) species to the western flats calls for a revision of Weimarck S
preliminary interpretation. The combination of an evident concentration of en-
demic taxa and intensive disturbance by man makes the western coastal flats a
priority area for research towards plant conservation. Williams (1972), referring to
Leucadendron floridum, stated: ‘“There is no doubt that, unless this historic
species is rigorously preserved, it will be extinct before long. It has already been
58 Journal of South African Botany
eliminated everywhere except in the Cape Point Nature Reserve, . . . where it is
now being rapidly overwhelmed by the introduced Australian Port Jackson wattle
[Acacia saligna Wendl., formerly A. cyanophylla Lindl.].’’ Leucadendron levi-
sanus, another flats endemic, was formerly “‘abundant’’ near Cape Town: ‘‘urban
development and the spread of the introduced Acacia cyanophylla Lindl., have
however virtually eliminated it from the Peninsula and it is fast approaching
extinction’ (Williams, 1972).
METHODS
Six study species in the family Proteaceae were chosen for the following
characteristics:
(1) endemism to a small continuous geographical range (maximum 700 km?)
within the western coastal flats, arbitrarily defined (Milewski & Esterhuysen,
1977) as from False Bay northwards, west of the Hottentots-Holland-Winterhoek
mountain chain to § 32 ° 30’ and eastwards along the intermontane flats to
Worcester, between sea level and 600 m altitude, and
small size and deteriorating conservation status of remaining total species-
population owing to man-made environmental changes (number of known
extant plants of each study species less than | 000).
(2
7
The study species, all evergreen perennials, were:
(i) Serruria ciliata R. Br.,
(ii) Leucadendron verticillatum (Thunb.) Meisner,
(111) Leucadendron flexuosum Williams,
(iv) Leucadendron floridum R. Br.,
(v) Leucadendron levisanus (L.) Berg., and
(vi) Diastella buekii (Gandoger) Rourke.
Methods were essentially similar to those used by Milewski (1977). Severe
limitations on the thoroughness of the investigation were imposed by the study
species’ relict status. All known remaining populations of each study species were
investigated, ‘“‘populations’’ of a species being arbitrarily defined (Milewski &
Esterhuysen, 1977) as stands farther than 2 km apart. Data were recorded from two
populations of all study species except Diastella buekii (until recently thought
extinct [Rourke, 1976]; a single rediscovered population was investigated) and
Leucadendron verticillatum (3 populations investigated). The physiognomy and
general floristics of associated vegetation were subjectively recorded for each
population.
Indirect (floristic) information was mainly used in comparing the habitats of
the study species. The four Leucadendron species grow as 1,0-2,0 m high,
densely branched, erect woody shrubs arising from a single stem; the two Serruria
species, forming 0,5-1,0 m high shrubs, have similar growth-form. Diastella
Habitat of Threatened Proteaceae 59
buekii grows as a dense “‘mat’’ (diameter 1,0-2,0 m) of creeping soft-woody
stems, originating from a single base at ground level and rooting adventitiously.
All species in the Proteaceae and Restionaceae growing within 0,5 m of plants of
each study species were recorded, data thereby being collected only within the
edaphic environment of the study species (plants of the study species cover a
larger area than do the herbaceous Restionaceous species studied by Milewski,
1977, necessitating a greater radius of investigation).
The search for members of these two families was continued until the list of
associates for each population appeared complete, i.e. until not more than one new
associate species was found with ten successively visited study plants, or until a
minimum of 20 plants, scattered throughout the population, had been investigated.
Lists, according to presence or absence, of associate species for each population
were combined to show the “‘between-population frequency’’ of each associate for
each study species. A list from one population of Serruria ciliata (apparently var.
congesta) was not included in the associate data, since the associated vegetation
appeared very different from that in which the typical form of the species grew,
and owing to the undecided taxonomic status of S. ciliata var. congesta at the time
of writing (Rourke, pers. comm.).
RESULTS AND DISCUSSION
1. General account of habitats
Serruria ciliata was restricted to deep, pale Tertiary sand on the northern Cape
Flats (Rourke, in prep.). The associated vegetation was a mainly restioid-ericoid
(sensu Taylor, in press) Coastal Fynbos, 1,0-1,5 m high. The species occurred
near Bellville and Bottelary as scattered plants with e.g. Berzelia abrotanoides,
Staavia adiata, Cliffortia polygonifolia and Passerina vulgaris in stands similar
to those associated with Chondropetalum acockii and Restio micans (Milewski &
Esterhuysen, 1977). Serruria ciliata var. congesta grew near Faure in vegetation
different in appearance, resembling Taylor’s (1972b) Thamnochortus-Passerina
Fynbos of Reddish Sands, with the ericoid shrub Passerina vulgaris. the tufted
| testioid Thamnochortus erectus, the low spreading shrub Myrica quercifolia and
several members of the Proteaceae and Restionaceae (Appendix 1). |
Leucadendron verticillatum was restricted to ‘‘fairly level’’ substrates “with a
clay subsoil’’ in ‘‘a small area where the divisions of Bellville, Paarl, Malmesbury
and Stellenbosch adjoin each other’’ (Williams, 1972). The associated vegetation
was an ericoid-restioid form of Coastal Rhenosterbosveld (sensu Taylor, in press).
The species occurred on the slopes of the Joostenberg as scattered emergent plants
in an open stand (canopy cover 40-50 %) comprising a main 0,7-0,9 m stratum
dominated by the ‘‘heath-like’’ shrub Elytropappus rhinocerous, and a ground
_ Stratum of wiry restioids (e.g. Restio cuspidatus) and forbs. Leucadendron verticil-
\latum occurred near Hercules Pillar as 2,5-3,0 m emergent shrubs in similar
60 Journal of South African Botany
vegetation, with 1,0-1,5 m proteoid (Leucadendron lanigerum), broad-leaved (e.g.
Olea africana) and ericoid (e.g. Eriocephalus africanus) shrubs.
Leucadendron flexuosum occurred only on “‘level alluvial soil containing a
large percentage of waterworn stones’’ in a small area of ‘‘the Breede River
Valley near Worcester’? (Williams, 1972). The associated vegetation was an
ericoid-restioid dry fynbos. The species grew near Hartebeest River Bridge as
scattered plants forming a 1,8 m emergent stratum in an open stand (canopy cover
40-50 %) comprising a main 0,3-0,4 m stratum of ericoid and other small-leaved
shrubs (e.g. Elytropappus gnaphaloides, Eriocephalus africanus, Relhania
squarrosa, Cliffortia ruscifolia) and wiry restioids (e.g. Restio cuspidatus).
Another population of Leucadendron flexuosum near Hartebeest River Bridge
occurred as groups of plants forming, with shrubs of Dodonaea viscosa, a 1,0 m
stratum emergent from a main stratum of tufted restioids (e.g. Thamnochortus
sporadicus) and ericoid shrubs.
Leucadendron floridum was restricted to sandy soil on the Cape Flats and
valleys of the Cape Peninsula (Williams, 1972). The associated vegetation was a
mainly graminoid-ericoid (sensu Taylor, in press) fynbos intermediate in appear-
ance between Coastal Fynbos and Mountain Fynbos. The species occurred near
Kommetjie and between Scarborough and Klaasjagersberg in dense stands (canopy
cover 80-90 %), comprising 1,0-2,0 m high tussock restioids (e.g. Restio com-
pressus) and other graminoids (e.g. Neesenbeckia punctoria), and ericoid and
other shrubs (e.g. Berzelia abrotanoides, Erica spp., Psoralea aphylla), and a
poorly defined lower stratum of sprawling soft shrubs (e.g. Cliffortia subsetacea)
and tufted restioids (e.g. Elegia fistulosa). This vegetation appeared similar to
Taylor's (1969) Berzelia-Osmitopsis Seepage Scrub Association and Restiona-
ceous Tussock Marsh Association.
Leucadendron levisanus occurred only “‘on sandy soil. . . on the Cape Flats
from the Fish Hoek valley north to the Koeberg’’ (Williams, 1972). The species
formed a scattered emergent stratum of 1,0-1,5 m shrubs in restioid-ericoid
vegetation similar to that associated with Serruria ciliata. Leucadendron levisanus
has previously been recorded growing with other emergents, e.g. Thamnochortus
erectus and Passerina vulgaris, among plants of e.g. Aspalathus hispida, Carpo-
brotus acinaciformis, Crassula cymosa, Psoralea fruticans, Rhus laevigata and
Ehrharta villosa (Taylor, 1972b).
Diastella buekii was restricted to “‘sandy situations’, on “‘the floor of the
upper Berg River valley’’ (Rourke, 1976). The associated vegetation was a mainly
restioid-ericoid fynbos intermediate in appearance between Coastal Fynbos
and Mountain Fynbos. The species occurred near Wemmershoek as scattered
plants in an open stand (canopy cover 50-70%) comprising a main 0,6—-0,7 m
stratum of tufted restioids (e.g. Thamnochortus sporadicus, Chondropetalum
nudum, Staberoha cernua) and ericoid shrubs (e.g. Cliffortia juniperina, Lachno-
spermum fasciculatum, Euryops abrotanifolius, Lachnaea capitata, Phylica stipu-
Habitat of Threatened Proteaceae 61
TABLE |
Restionaceous and Proteaceous associate species, according to “‘between-population fre-
quency’’, of six Proteaceous study species: 1 = Leucadendron floridum, 2 = Leucadendron
levisanus, 3 = Serruria ciliata, 4 = Diastella buekii, 5 = Leucadendron flexuosum, 6 =
Leucadendron verticillatum. Numerator is the number of populations in which the associate
was recorded within 0,5 m of plants of each study species; denominator is the total number
of investigated populations of each study species; **+’’ indicates associates recorded in the
same stand of vegetation as the study species but not within 0,5 m.
Occurrence with study species
Associate species 1 2 3 4 ] 5 [ 6
lege Silage VAUUN oo ogocessos eaaneocooooE /2
RGSTORGOMPLESSUSMIN OUD rete tate le rire ele ei 2/2
Diastella divaricata (Berg.) Rourke ............. 1/2
Leucadendron laureolum (Lam.) Fourcade ....... 1/2
R@STG CEG ITENS, occoconcm one géosenosoodsDd 1/2
[El eanG: GLITCH WIENS caccsecoedenodnosoudEDT 1/2
Elegia asperiflora (Nees) Kunth ................ Pyf), |) ar
Senjunanelomeratal (I2.) ROBIN. 2-22.22 o3--- AYfP2 \\ WYP
Restouetmagonus Phunbie =... 2-5. --2 2-6: Hp || YP
SGI: CHIT) VE 6 oagabece caneogengganuodd 1/2
RCSD TALUCATE NOt Geacecaudscuaduucboodcod 1/2
IEWARG! WAR AgaEE WES Geonebcoess0b0edodeoeRd +
Bilecraiprominens bilians ee. ee i +
RGSRO SHORTS VANEISS coccocobocecuouocucudn +
Protea scolymocephala Reich. ..........------- + +
iResttoymicans (Kunth) Nees =--.-.-.-.--------- V2 |) Wy
Thamnochortus punctatus Pillans .............-. 1/22
Diastella proteoides (L.) Druce .......----.---+ + | 1/2
Leptocarpus impolitus (Kunth) Pillans ........... 2/2
Leucadendron levisanus (L.) Berg. ......-----++-: 1/2
Leucadendron cinereum (Solander ex Ait) R.Br.... 1/2
Chondropetalum acockii Pillans .........------- 1/2
Thamnochortus obtusus Pillans ...........--+++: 1/2
Prataa: juries VSGOUE cnc pocedoneecuauaccooe4 1/2
Staberoha distachya (Rottb.) Kunth ............. 1/2
PrGae (GIG WL) boo o65eceabouuooden peeoouI4 ale
Nenuria furcellata RBI... . 2222626602 - 9-20" +
Leucospermum hypophyllocarpodendron (L.)
DIGS: pe cewietje oe RO ein Oc en eae ype, \\ AUP
Hypodiscus aristatus (Thunb.) Nees......-------
Restio curviramis Kunth ...........-----++++::
EIOR OTITIS EOI sec e-.. 2-5. sere eee: 1/1
Restio pedicellatus Mast.......-..-++--+++:++++:
Restio monanthus Mast.........-----++---+°+::
Resttonwalliehiwt Mast) seeecie.--- 26-227 4° ss
Thamnochortus sporadicus Pillans .......-------
EHO Gu, SION, MW cooenancu. cop den oeeooe mapaiag 5/2
Cannomois acuminata (Thunb.) Pillans .......---
Leucospermum calligerum (Salisb. ex Knight) 1/2
POUTTG onc oc Good Gels Hoe Ebon uOROoem sonore
Leucadendron brunioides Meisn. ....-.-.---+++:
62 Journal of South African Botany
TABLE | (contd)
Occurrence with study species
Associate species
Leucadendron chamelaea (Lam.) Williams .......
Thamnochortus bachmannii Mast. ..........----
IRA CHT HATE JAWENG occo0c0esc00000000000006
Leucadendron lanigerum Buek ex Meisn. ........ 2/3
Elegia squamosa Mast. ...........-.+--+------ 2/3
Hypodiscus paludosus Pillans ...........------- 2/3
Elegia sp. nov. Esterhuysen 34411.............. 2/3
ROSH CAPLORIS ROMO, 2 o0000000000000000000008 2/3
Praga @dopaia WUD, coccaceccc0000000000000 1/3
IROSHO GEORAGWIGIETILOS INES 6 o5000000000000000006 Dp) \\ WP |) Wy
Restio) bifurcus Nees ex Mast.......--.--....-.- W2 || 2 || Wye
Chondropetalum nudum (Nees) Rottb. .........-- DP \|\ DP = 1/1
JABHO WORT BTUGTTO IMESG 6 9.0 00000000000000000008 2/2 ae lil
Elegia neesii (Mast.) Mast. ..............------ 2/2 1/1
Elegia coleura Nees ex Mast.............+---++- 1/2 1/2
Chondropetalum tectorum (L.) Pillans ........... 1/2 | 1/2
Willdenowiarsulcatas Mas taapencnce eee de) BYP | iyi
Willdenowiarhumiliss Masten nec crc + 1/2 | 1/1
Hypodiscus willdenowia (Nees) Mast. ..........- aF 2 1 iil
Staberoha cernua (L.f.) Dur & Schinz........... 2/2 oF W2 | iil || Dp
Restiojpaludosusmrilansineeenooe cect errr WW |) Wf |) se Wil | ye
Leucadendron salignum Berg.............+.++-- + V2) Wil |) aye
(HARTA! OCIIIPIORE, VAUININ 3 0000000000 00005d00000C W2 || QQ | Wye 1/2 | 1/3
IROSHO Fs WOo 2 9 000000000000000000000000000 + 1/2
Willdenowia arescens Kunth................... 2/2 1/2
ILGDOCTDS IBOTOUS IES, 2 o0c0eb00000000000C 1/2 W2 | QB
Chondropetalum rectum (Mast.) Pillans .......... 1/2 2/3
Senna. loxmmiaarall IRB, oo 00 cee c000a08e00Ke000 + +
Leptocarpus vimineus (Rottb.) Pillans ........... aP
ROS CU DICGHIS WNIT, oo000o0000000000000004 + Wil || Be | Bye
+
+
RESTO SD WN 3B ccccaocacceesoooncovcnvesccr
Thamnochortus fruticosus Berg. ..............--
Wilidenowiakstriataalhunb yee eee eneeeie
laris). Scattered proteoid, ericoid and other shrubs (e.g. Leucadendron salignum,
Passerina vulgaris, Berzelia abrotanoides, Anthospermum aethiopicum, Rhus
angustifolia, Diospyros glabra) emerged to 1,2 m.
2. Habitat similarity
Table 1 shows sets of Proteaceous and Restionaceous associates for the study
species. The column sequence in Table 1 interrelates the study species on the basis
of associates shared, owing to the arrangement of rows and columns. Table 2
shows percentage similarity between these sets.
Similarity values were generally low; the only pairs of species showing more
than 30 % similarity between sets of associates were Leucadendron floridum with
Habitat of Threatened Proteaceae 63
f TABLE 2
Percentage similarity between sets of associates for six study species of Proteaceae, using
“‘between-population frequency’’ data on Proteaceous and Restionaceous associates (Table
1) to calculate the Czekanowski coefficient. The formula used was es x 100 (Bray &
Curtis, 1957), where w = number of associate species in common, A = number of
associates for one study species, and B = number of associates for another study species.
Study species
1
eee cucadendronponidumeRo Bia --rie scenes le
. Leucadendron levisanus (L.) Berg ...........
MNS CLLURIAKCHIQIQURG BT geri. ues ta ete melons
. Diastella buekii (Gandoger) Rourke ..........
. Leucadendron flexuosum Williams ...........
. Leucadendron verticillatum (Thunb.) Meisn. ...
DANnhWNe
L. levisanus, and Diastella buekii with Leucadendron flexuosum. Leucadendron
verticillatum and L. flexuosum had sets of associates largely different (17,4 %
similar) from each other and from those for the other two congeneric study species
(3,9-20,0 % similar). The set of associates for Serruria ciliata was more similar to
that for L. levisanus (one of its associates) than to those for any other study
species.
This numerical assessment of similarities can be viewed as a reflection of
habitat similarity for the study species, based on the assumptions set out in the
introduction. The habitats of the study species thus appear in general to be distinct
from each other.
3. Discussion
Leucadendron levisanus, L. floridum and Serruria ciliata all formerly occurred
in partial sympatry on the northern Cape Flats. Each of the remaining three study
species was allopatric with all other study species.
Leucadendron levisanus and L. floridum, largely sympatric, have been re-
corded from the same localities (Williams, 1972). A difference in their habitats
has, however, been hinted at in the literature: L. /evisanus has been recorded from
‘‘sandy soil, often very damp in winter’, while L. floridum *‘is always found
growing near to streams or in damp places . . . it will grow with its roots
submerged for long periods in waterlogged ground’’ (Williams, 1972). The chief
habitat difference between these two congeners thus appears to be their different,
although possibly overlapping, tolerance ranges with regard to soil moisture.
Serruria ciliata evidently also has a discrete association with relatively poorly
drained substrates, ranging from those supporting L. levisanus to better drained
sands supporting the very habitat-specific Chondropetalum acocki (the habitats of
64 Journal of South African Botany
Chondropetalum acockii and Restio micans, associates respectively of S. ciliata
and L. levisanus, have been described by Milewski & Esterhuysen, 1977).
The habitat of Diastella buekii, as expressed by its set of associates, is more
similar to those for L. Jevisanus and S. ciliata than to that for L. verticillatum; the
habitat of Leucadendron flexuosum appears equally similar to these two groups.
The sand associated with D. buekii, probably of transported Table Mountain
Sandstone origin, is apparently that of an intermontane alluvial plain, as is the
substrate associated with L. flexuosum. However, D. buekii receives more than
twice the average annual precipitation (800-900 mm, localities given by Rourke,
1976) received by L. flexuosum (300-400 mm, localities given by Williams,
1972), which grows on the edge of the Karoo.
The interrelationships of the habitats of the study species, thus interpreted from
sets of associates, agree with the author’s field impressions. The evident interrela-
tionships of the vegetation associated with the study species, assessed subjectively
on the basis of physiognomy and general floristics, also support these conclusions.
The main variation in the vegetation of the western flats appears to be in
response to two edaphic gradients, the first determined by the proportions of sand
and clay in the soil and the second by site drainage (e.g. Talbot, 1971; Taylor,
1969, 1972b, in press). The sand-clay ratio evidently results, at the extremes of the
range, in the basic dichotomy outlined by Acocks (1975) and Taylor (in press),
that of Coastal Fynbos (Coastal Macchia) on deep pale Tertiary sand and Coastal
Rhenosterbosveld on clay-rich soil weathered from Malmesbury shale. Leucaden-
dron levisanus, L. floridum and Serruria ciliata are species of the sand biotope,
while Leucadendron verticillatum is a species of the clay biotope. Leucadendron
flexuosum can perhaps be regarded as a species of an intermediate, relatively dry
environment; Diastella buekii can perhaps be regarded as a species of a sandy
environment intermediate between the Coastal Fynbos and Mountain Fynbos (i.e.
Acocks’ Macchia on Table Mountain Sandstone) biotopes.
The apparent dissimilarity between the habitats of the study species, including
those which grow in geographical proximity in apparently similar environments,
reflects the complex floristic pattern within Coastal Fynbos and Coastal Rhenoster-
bosveld. Preservation of as wide a range as possible of coastal flats endemics and
the plant communities in which they occur calls for the strict preservation of all
remaining relics of natural vegetation until a detailed phytocenological survey can
provide the basis for optimal placing of permanent flora reserves.
ACKNOWLEDGEMENTS
I wish to thank Professor E. A. C. L. E. Schelpe for the use of facilities at the
Bolus Herbarium, University of Cape Town. Dr A. V. Hall, project leader of the
Rare and Endangered Plant Species Research Group, supervised the study. Miss E.
Esterhuysen and Dr J. P. Rourke gave freely of their expertise, and assisted with
identification of Restionaceae and Proteaceae respectively. Dr E. J. Moll, Dr P.
Habitat of Threatened Proteaceae 65
Bridgewater and Mr H. P. Linder commented on the manuscript. Financial
assistance provided under the National Programme for Environmental Sciences is
gratefully acknowledged. I am indebted to Mrs M. L. Jarman and Miss C.
Davidge for typing the manuscript.
REFERENCES
Acocks, J. P. H., 1975. Veld types of South Africa. Mem. bot. Surv. S. Afr. 40.
BoucHEerR, C. 1972. The vegetation of the Cape Hangklip area. M.Sc. thesis.
Cape Town: University of Cape Town.
Bray, R. J. and Curtis, T. T., 1957. An ordination of upland forest communities
of southern Wisconsin. Ecol. Monogr. 22: 325-349.
MiLewski, A. V. and EsTERHUYSEN, E., 1977. Habitat of Restionaceae endemic
to western Cape coastal flats. J] S. Afr. Bot. 43 (4): 233-241.
Mitewski, A. V., 1977. Habitat of Restionaceae endemic to the south-western
Cape coastal flats. J/ S. Afr. Bot. 43 (4): 243-261.
RARE AND ENDANGERED SPECIES RESEARCH Group. Unpublished reports on
distribution, habitat and status of threatened plant taxa in the south-western Cape,
1976. Cape Town: Bolus Herbarium, University of Cape Town.
Rourke, J. P., 1972. Taxonomic studies on Leucospermum R.Br. Jl S. Afr. Bot.
Suppl. vol. 8.
RourkKE, J. P., 1976. A revision of Diastella (Proteaceae). JI S. Afr. Bot. 42
(3): 185-211.
Rourke, J. P., in preparation. A revision of the genus Protea L.
Rourke, J. P., in preparation. A revision of the genus Serruria Salisb.
TatBot, W. J., 1971. South western Cape Province. The South African land-
scape: 1. South African Geographical Society.
Taytor, H. C., 1969. Vegetation of the Cape of Good Hope Nature Reserve.
M.Sc. thesis. Cape Town: University of Cape Town.
TayLor, H. C., 1972a. Fynbos. Veld & Flora 2: 68-75.
Taytor, H. C., 1972b. Notes on the vegetation of the Cape Flats. Bothalia 10
(4): 637-646.
Taytor, H. C., in press. Phytogeography and ecology in Capensis. In: M. J. A.
Werger (ed.), Biogeography and ecology in Southern Africa. The Hague: Junk.
Wermarck, H., 1941. Phytogeographical groups, centres and intervals within the
Cape Flora. Acta Univ. lund. N.F. Avd. 2, 37 (5).
Wuittaker, R. H., Levin, S. A. and Root, R. B., 1973. Niche, habitat and
ecotope. Am. Nat. 107 (955): 321-338.
Wituiams, I. J. M., 1972. A revision of the genus Leucadendron (Proteaceae).
Contr. Bol. Herb. 3.
Appendix 1: Restionaceous and Proteaceous associates of Serruria ciliata var. congesta
(100 plants investigated in 1 population).
Leucadendron salignum, Protea scolymocephala, Serruria burmannit, Hi eae i
erectus, Thamnochortus fruticosus, Thamnochortus obtusus, Hypodiscus wt ae 9
gia vaginulata, Willdenowia striata, Chondropetalum nudum and Restio cuspidatus.
JIS. Afr. Bot. 44 (1): 67-81 (1978)
STUDIES IN THE GENUS CASSIA IN SOUTH AFRICA: 2. NOTES ON
CASSIA ITALICA (MILL.) LAM. EX F. W. ANDR.
K. D. GORDON-GRAY
(Bews Botanical Laboratories, University of Natal, Pietermaritzburg)
ABSTRACT
Two characters (stigma form and ovary indumentum) previously unused in subspecific
delimitation within Cassia italica (Mill.) Lam. ex F. W. Andr. are considered and additions
are made to features already useful in infraspecific classification in this taxon. Two
subspecies, the one comprising three forms, are confirmed for the species in southern Africa
a Cae ranges for the taxa are given. Some previously perplexing variants are
clarified.
UITTREKSEL
’N STUDIE VAN DIE GENUS CASSIA IN SUID-AFRIKA:
2. AANTEKENINGE OOR CASSIA ITALICA (MILL.) LAM. EX F. W. ANDR.
Twee kenmerke (stempelvorm en ovariumbeharing) wat voorheen nie in die afbakening
op subspecies-vlak in Cassia italica (Mill.) Lam. ex F. W. Andr. gebruik is nie, word in
oorweging geneem en word tot kenmerke wat alreeds nuttig is in die indeling op
infraspecies-vlak in hierdie takson gevoeg. Twee subspecies, een bestaande uit drie forms,
word bevestig vir die species in suidelike Afrika en die verspreidings vir die taksons word
aangegee. ‘n Paar variante waaroor daar voorheen onduidelikheid was, word toegelig.
Brenan (1958) outlined the wide geographical range of Cassia italica (Mill.)
Lam. ex F. W. Andr. and analysed its ‘‘. . . remarkable range of variation,
_..?. On leaf, inflorescence and flower characters, this author was able to
recognize three races which he appropriately designated subspecies because of
their mostly distinct distributions and because of occasional inconstancies in the
correlated characters by which they were distinguished. Table 1 gives a summary
of these distinctions and of the geographical ranges of the subspecies as deter-
mined by this author. The southern race, subsp. arachoides, is undoubtediy the
most variable. Much South African material corresponds with the holotype of this
subspecies (South Africa, Griqualand West, Burchell 1680, K), but Brenan (1958)
enumerated three variants that diverge from this type and he listed other “*. . .
perplexing specimens. . .”’.
In revising Cassia for the Flora of Southern Africa, it became necessary to
take up the study of this subspecies and of subsp. micrantha, both of which reach
the Flora area, and to pursue the ‘‘perplexities’’ that Brenan had so ably exposed.
This subsequent study has revealed the presence within C. italica of two
characters convenient in infraspecific diagnosis that were unused by Brenan.
Accepted for publication 19th July, 1977.
67
68 Journal of South African Botany
These are:
1. Stigmatic form
In subsp. arachoides the style is long and more or less coiled: it terminates in a
stigmatic zone only slightly wider than the style breadth which narrows again to
end in a small more or less circular opening (Fig. la, b: Fig. 2b, c).
In subsp. micrantha the style is visibly shorter and does not coil, but merely
recurves towards the ovary: the terminal stigmatic zone is clearly wider than the
style and asymmetrically trumpet shaped (better developed adaxially than abaxially)
with an opening as wide as the stigmatic expansion. In herbarium specimens the
delicate margin of this stigmatic tissue is usually reflexed, but this is not often the casein
living flowers (Fig. lc: Fig. 2a).
In the few specimens of subsp. italica I have examined, the stylar and
stigmatic form agreed precisely with that of subspecies arachoides. Probably this
could do with further checking.
Because style and stigma persist after pollination and can be seen clearly
exposed on young developing fruits, the form of these structures provides a useful
means by which to distinguish the subspecies micrantha and arachoides. The
criterion seems reliable (as is to be expected considering the use that has been
made of stigmatic form in aiding the delimitation of genera in papilionates, for
example Dolichos and its allies, Verdcourt, 1970), for among the more than 200
specimens examined, only one possible intermediate was encountered (see De
ovaries mm,
hairs 2.5mm
obttetatd
roan
LN OIG
BiGaale
CASSIA ITALICA, developing ovaries with their indumentum types: a. subsp. arachoides
minute-haired form with trichomes patent (left) and curved (right); b. subsp. arachoides
villous form; c. subsp. micrantha.
Genus Cassia in SA; Notes on Cassia italica 69
Winter & Leistner 5756 later) but this had clearly the stigmatic structure of subsp.
micrantha.
2. Indumentum of developing ovary and mature legume
Vesture, together with some consideration of the hair types comprising it, has
been used before in infraspecific classification within C. italica. Burtt Davy (1932)
established C. obovata var. pilosa for plants with stems, rhachises and leaves
pilose, while Brenan (1958) included indumentum among the characters he used
(see Table 1). No author yet seems to have commented upon the indumentum of,
or its absence from, young ovaries or mature fruits. This may be because the
trichome types present are those of the vegetative organs, but on the ovaries in
particular, the hairs are closely packed and thus readily observed, especially as the
outer floral parts fall away after fertilization to leave the young ovaries exposed.
This character aids particularly in the recognition of variants within subsp.
arachoides, but can be used as an additional means by which to distinguish the two
subspecies.
In the subspecies micrantha ovaries are densely clothed in curved, appressed,
white hairs which give them an almost hoary appearance (Figs Ic: 2f). Occa-
sionally some of these hairs are swollen and inversely bottle-shaped (perhaps
serving as water reservoirs?). As growth and maturation of the fruit takes place,
the hairs become spaced so that the mature valves are sparsely appressed-
pubescent.
In subsp. arachoides the indumentum is not uniform and four variants or forms
may be recognized:
Variant Ovary Legume Trichome type
(representative
of whole plant)
A (typical form) glabrous glabrous short, straight,
patent,
scattered (Fig.
3a, g)
B (minute-haired densely sparsely minute, curved
form) appressed- appressed- appressed (Figs.
pubescent pubescent la, right; 2d;
3d)
C (minute-haired densely sparsely minute, straight,
form) pubescent pubescent patent (Figs. la, left;
2e; 3f)
D (villous form) densely villous villous long, slender,
straight, patent,
fine hairs (Figs.
1b; 2g; 3h)
70 Journal of South African Botany
Fic. 2.
CASSIA ITALICA, scanning electron micrographs of: a. subsp. micrantha, stigma (X 105);
b. & c. subsp. arachoides, stigma (X 105); d. subsp. arachoides minute-haired form
(trichomes curved), ovary indumentum (X 300); e. ditto but trichomes patent (X 300); f.
subsp. micrantha, ovary indumentum (X 300); g. subsp. arachoides villous form, ovary
indumentum (X 135).
Genus Cassia in SA: Notes on Cassia italica
xs ved
ge ge “Rep
28 gn ee CGF 0 =
chee. ox
Fic. 3.
CASSIA ITALICA, scanning electron micrographs of: a. subsp. arachoides typical form,
leaf surface showing trichomes and papillae (X 125); b. ditto, papillae enlarged (X 200); e.
ditto, further enlarged, note stoma (X 800); g. ditto, single trichome enlarged (X 600); c.
subsp. micrantha, leaf surface showing single trichome (X 600); k. ditto, further enlarged
(X 1625); d. subsp. arachoides minute-haired form, leaf surface showing single curved
trichome (X 600); f. ditto showing two patent trichomes (X 600); h. subsp. arachoides
villous form, leaf surface showing base of trichome and several stomata (X 300).
71
U2 Journal of South African Botany
On the basis of stigma type and indumentum it has been possible to clarify one
of the doubts expressed by Brenan, 1958, and to provide a few more facts relevant
to the understanding of infraspecific variation in C. italica.
Burtt Davy 7040 (K) from hotel-yard at Pienaars River Station, Transvaal, that
was doubtfully determined by Brenan /.c. as subsp. micrantha, has been re-
examined by Dr J. H. Ross, SA Liaison Officer at Kew, whom I should like to
thank. This specimen has the stylar form typical of subsp. arachoides. It is
therefore in accord with distributional patterns of variants within C. italica
determined in the present study, for no specimens of subsp. micrantha have been
found further east than Lugard 201 from Kwebe Hills, Botswana, approximately
S 20° 40’; E 23° 5’ (also cited by Brenan, 1958).
The only other specimen seen that did not immediately fall within one or other
subspecies on the basis of the two characters mentioned above was De Winter &
Leistner 5756 from Otjinungua, north western South West Africa. This has the
stylar form of subsp. micrantha but the indumentum of subsp. arachoides, to
which taxon it was referred by Schreiber, 1967. (I prefer to place it with subsp.
micrantha as, in my opinion, it also agrees with this taxon in petiole length,
inflorescence/leaf ratio and leaflet apex). It is from an area where both these
subspecies are thought to be sympatric (but this must be checked on the ground, if
possible). Such a specimen suggests that outbreeding is possible between these
subspecies, as would be expected.
From study of the indumentum of ovary and fruit in the considerable amount
of material of subsp. arachoides available in herbaria, it has been possible to gain
better knowledge of, and roughly to delimit the areas occupied by, the variants
within this subspecies, some of which were listed by Brenan 1958: 243.
Variant A represents the typical form of subsp. arachoides. Here ovary and
fruit are glabrous. Plants are sometimes described as glabrous and glaucous. The
latter epithet is well applied, but strictly plants are not glabrous for magnification
reveals scattered short straight patent hairs on leaves and stems that Brenan
referred to as “‘Indumentum saepius breve, sparsum, patens. . .’’ (Fig. 3a). Often
on young stem apices in particular, and especially in plants growing where
environments are extreme, the hairs are interspersed with small, short-stalked or
subsessile elliptical glands that impart a certain viscidity. The leaves are also often
papillate as is clearly revealed by electron scanning (Fig. 3a, b, e). This typical
form is the “‘dryland”’ variant within subsp. arachoides, for plants occupy harsh,
sub-desertic environments in South West Africa, Botswana, the northern central
Cape Province, the Orange Free State and the western Transvaal (Fig. 4). A
majority of southern African plants belong with this form.
Variants B and C represent Brenan’s variant (2). These plants are green, not
glaucous, and again appear glabrous to the naked eye, but microscopic examina-
tion reveals minute hairs more closely packed than in the typical form. Ovaries
and fruits are hairy, not glabrous. In some plants (variant B) the hairs are curved
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CASSIA ITALICA,
74 Journal of South African Botany
and appressed; in other plants (variant C) the hairs are straight and patent. Brenan
did not differentiate between the two hair types, so it is uncertain whether he
noticed them or not. Of the specimens he cites I have not seen Trapnell 744 from
Natal, but Barnard & Mogg 966 and Codd & De Winter 5009, both from the
Transvaal, have curved hairs. Natal and Swaziland plants more commonly have
straight, patent hairs, while those from the Transvaal are often curved, but this is
not absolute.
These minute hairs, be they curved or straight, are distinguishable under
magnification ( + X 20) from the hairs of subsp. micrantha and from hairs of the
typical form of subsp. arachoides. Especially on the ovaries is this the case, for
the longer appressed hairs of subsp. micrantha overlap to a degree that the minute
hairs do not (compare Fig. 2d, e, f). This minute-haired form is the “‘moist-land”’
variant within subsp. arachoides. This appellation is, nevertheless, something of a
misnomer, for plants, like those of the typical form, also occur in open, dry
habitats, but such conditions in eastern Southern Africa are far less harsh than are
the sub-desertic conditions of the west. The minute-haired form occurs in Rhode-
sia, Mogambique, Swaziland, the Transvaal and Natal as far south as the Tugela
Valley. In the Transvaal it is mainly central and eastern: towards the west there is
overlap with the typical form.
Variant D represents Brenan’s (1958: 242) variant (1) and, as this author
stated, is also probably the form that Burtt Davy (1932: 325, Type: Rogers 22504,
Transvaal, Pilgrims Rest distr., Newington—not seen by me) established as C.
obovata var. pilosa (non Senna obovata (Collad.) Batka var. pilosa Batka which is
Cassia italica subsp. italica). Also probably belonging here is Moss & Rogers 36,
which Baker (1930) and following him Burtt Davy (1932) tentatively placed with
C. holosericea Fresen., which is a north-east African species with villous pubes-
cence resembling that of Variant D, but with smaller flowers and pods that lack a
line of crests.
Variant D is the villous form within subsp. arachoides. The hairs to the ovary,
mature legume, peduncle and the vegetative organs are long, fine, straight, patent
and closely packed so that the indumentum is villous. Brenan cited only three
specimens of this form, but the present survey has shown that it is well established
in the Transvaal north of S 25° and between E 28°—30° 30’. Rogers 6074 from
Francistown, Botswana is not quite as evidently hairy to the naked eye as are most
specimens referred to this form. If this gathering and Rogers 22504 are included in
variant D, then its distributional limits are extended slightly east and west.
Other characters correlated with variation within C. italica
3. Leaflet apex
Burtt-Davy (1932: 325) established C. obovata var. mucronata (Type: Galpin
750, Transvaal, Barberton, near Queen’s River) for plants in which the leaflets are
drawn out into a mucron 0,5 to 1,0 mm long. It is not necessary to maintain this
——
Genus Cassia in SA: Notes on Cassia italica 75
taxon, for this character appears to be correlated with the development of minute
hairs, so that it is a feature of the minute-haired form of subspecies arachoides. It
is not limited to this entity, however, because plants of subsp. micrantha usually
develop leaflets with similar mucronate apices. In contrast, the typical form of
subsp. arachoides generally has emarginate leaflet apices. The only deviation from
this is in occasional plants where whole leaves (and their leaflets) are very much
larger than usual and are assumed to have developed during exceptionally
favourable growth periods. The apices to these larger leaflets are rounded rather
than emarginate, sometimes with a poorly-defined mucron. In the villous form of
subsp. arachoides apices vary from more or less emarginate, to rounded, some-
times with a poorly-defined or short mucron.
4. Legume crests and legume shape
In C. italica, each valve of the legume develops a lengthwise midline of
discrete, small evaginations called ‘‘crests’’. Burtt-Davy (1932: 325) described
Galpin 750 (type of his var. mucronata) as bearing pods lacking crests, but on the
several sheets of this gathering in various herbaria, some pods show some
evidence of evaginations. Study has shown that it is rather exceptional to find all
legumes developed by a plant without trace of crests but sometimes they are
scarcely discernible as in the (straight) minute-haired form in Natal (Ward 3613,
PRE, NH, NU). Small, poorly-defined crests are known only among plants from
eastern southern Africa: thus this feature seems correlated with the minute-haired
form, but it is by no means always developed here, however, and as Brenan (1958:
243) suggested, seems no more than a chance variation.
Legume shape also shows variation. This seems to correlate in part at least
with the taxa already recognized within C. italica. Pods of plants of the typical
form of subsp. arachoides are usually more or less sub-orbicular with glabrous
valves on which the veins are clearly marked (Fig. 4), whereas pods of all other
variants including subsp. micrantha develop longer, narrower, more or less oblong
pods, in which the valves carry an indumentum and where the veins are usually
less obvious (Fig. 4). It is tempting to think these differences in shape are the
result of differences in rates of growth concomitant with the differing environ-
ments under which the plants grow, but this is not supported in the case of fruits
of subsp. micrantha.
5. Petiole length
Reference to Table 1 shows that Brenan (1958) recognised subsp. arachoides
as having shorter petioles than either of the other two subspecies. Thus he found
perplexing certain specimens which, while agreeing with other characters of this
taxon, yet possessed long petioles. Some of these specimens otherwise resembled
the holotype of subsp. arachoides; others belonged with the minute-haired variant.
In attempting an explanation for these long petioles, Brenan considered that they
76 Journal of South African Botany
might be produced on young actively growing shoots, or that they might result
from introgression of subsp. micrantha into subsp. arachoides.
During the present study a number of points have become clear:
(i) within the distributional range of the typical form of subsp. arachoides it is
not unusual for short petioled leaves to be succeeded on the same stem by
leaves with much longer petioles (Werger 235, Orange Free State, Hopetown
distr. near Luckhoff, has leaves with petioles ranging roughly in sequence
from 5,0-15 mm (Fig. 5)). Also in occasional localities, plants with petioles
up to 19 mm long have been repeatedly collected. These localities mostly lie
between S 26°-30° and E 23°-26° and include the Herbert district and
Bloemhof from where, respectively, Burchell 1764 and Leistner 53 (both
cited by Brenan, 1958: 243) were obtained. Other localities and specimens
worth mentioning are: Vryburg, Armoedsvlakte, Sharpe 7056; Bloemhof,
Kameelpan, Christiana, Theron 427; Prieska, Bryant J178. Then, too, in other
dry areas such as South West Africa, a single gathering may be much more
robust and larger in all its vegetative parts than other specimens from the
same general locality ([hlenveldt 1888, South West Africa, distr. Rehoboth,
farm Bergland).
(ii) Within the minute-haired form it is usual for leaves to have petioles longer
than the range given by Brenan (1958) for subsp. arachoides (see Table 1).
Apart from the specimens of this group cited by Brenan as “‘perplexing’’,
there are many others that exhibit the same feature (Edwards 3189, Natal,
Ngotshe distr., 142 ml. from Pongola bridge on Magudu road, carries leaves
with petioles from 10-18 mm long on the same plant).
(iii) Considering the range in petiole length recorded for leaves on single stems, it
seems probable that within the typical form of subsp. arachoides at least, this
is a seasonal, or a habitat, phenomenon associated with favourable growing
conditions. This is surely not unexpected considering the response that
usually results from the presence of moisture in inhospitable environments.
On the basis of present distributions it seems unlikely that introgression from
subsp. micrantha is the explanation underlying these longer petioles since
most specimens are known from near the south eastern distributional limit of
the typical form. But what is the explanation underlying the presence of the
villous form of subsp. arachoides in the Pietersburg area of the Transvaal and
probably further north? Is this a case of introgression, but with a taxon not
now represented there?
6. Relative lengths of leaves and inflorescences and flower size
Brenan, 1958, collectively used three characters to classify C. italica at
infraspecific level. These were:
Genus Cassia in SA: Notes on Cassia italica 77
Arent), HERBARIUM PRETORIA
Hey
FIG, 5.
CASSIA ITALICA subsp. arachoides typical form: Werger 235 from near Luckhoff, stem
showing variation in leaf and leaflet size.
78 Journal of South African Botany
TABLE 1.
Characters used by Brenan (1958) to differentiate subspecies within C. italica
subsp. italica subsp. micrantha subsp. arachoides
petiole length mostly 10-35 mm mostly 10-25 mm mostly 3-12 mm
raceme length 60-230 mm 20-80 mm 70-220 mm
raceme length: mostly raceme ex- raceme shorter mostly raceme ex-
subtending leaf ceeding subtending | than subtending ceeding subtending
length leaf, sometimes leaf leaf
subequal
sepal length 8,0-13,0 mm 5,0-8,0 mm 8,0 mm
petal length 9,0—20,0 mm 8,5—9,0 mm 10,0-12,0 mm
petal width 5,0-10,0 mm 3,5—4,5 mm 5,0-6,0 mm
anther lengths
longest 8,0-14,0 mm 5,5-6,0 mm 8,0 mm
medium 4,0-6,0 mm 2,5-3,5 mm 2,5-3,0 mm
shortest 1,5-2,5 mm 1,25 mm 1,0 mm
indumentum appressed-puber- appressed- often short, sparse,
ulous, occasionally | puberulous spreading, rarely
shortly spreading- dense, or appressed-
pubescent puberulous
distribution Cape Verde Is., Ethiopia, British Rhodesia,
Senegambia & Somaliland, Uganda,| Mocgambique,
Rio de Oro to Kenya, Tanzania, South West Africa,
Egypt & Somali- Botswana & South | Botswana, Swaziland
land & south to West Africa. & South Africa.
Nigeria, the Sudan | Also most of India.
& Ethiopia. Also
Israel, Arabia,
Persia, Baluchistan
& n.w. India.
(a) ratio of subtending leaf length to inflorescence length
(b) petiole length
(c) flower size
He was careful to stipulate that none of these characters considered separately
was absolute in enabling distinction to be made among the three subspecies,
italica, micrantha and arachoides. The present work has shown that when these
criteria are used (and they are helpful) the following additional attributes will
make them easier to apply and more reliable in differentiation:
(i) inflorescence length for comparison against subtending leaf length: ideally
measurement should take place as the first flower opens.
Genus Cassia in SA: Notes on Cassia italica
TABLE 2.
Main features useful in differentiating subspecies and forms within Cassia italica
in southern Africa.
79
subspecies arachoides micrantha
forms typical minute-haired villous
stigma trumpet-shaped contracted into an aperture narrower than
(Fig. 2a) style (Fig. 2b, c)
style curved, +
3-4 mm more or less circinnate, 6-7 mm long (Fig.
long (Fig. Ic) la, b)
ovary
indumentum hoary lacking hoary villous
ovary
trichomes see Fig. Ic; lacking see Fig. la; see Fig. 1b;
2f Dane 2g
leaf
trichomes microscopic short, patent microscopic villous
appressed sparse (almost | straight or
microscopic) curved
petiole
length in
mm 9-23 (—25) (3-)6-12(-19) | 8-15-20) (3-)5-12
leaflet apex mucronate emarginate to mucronate emarginate to
rounded rounded
petal length
in mm 5-7(-9) 7,0-12,0 7,0-12,0 10,0-18,0
pods + oblong + suborbicular | + oblong + oblong
trichomes short,| glabrous, veins trichomes villous
appressed clearly marked | minute, crests
scattered sometimes
+ lacking
distribution S.W. Africa; S.W. Africa; Swaziland; N. Transvaal;
Botswana Botswana; central & E. Botswana
N. Cape; W. to | Transvaal,
central Trans- | N. Natal &
vaal Tugela Basin
(ii) petiole length and flower sizes:
attribute separately for each form or variant wl
collectively for subsp. arachoides.
These additions are given in Table
infraspecific variation within C. italica as this is present
it is necessary to give a range for each
thin this subspecies and not
2 which may serve as a summary for
ly known in South West
80 Journal of South African Botany
Africa, Swaziland and South Africa. A key permitting identification of the
infraspecific taxa within C. italica represented in these countries is provided.
The following conclusions may be drawn:
1. No plants with all the size dimensions of subsp. italica have been seen from
the area covered by the Flora of Southern Africa. There is no doubt, however,
that the subsp. italica and arachoides are closely allied and differ quantita-
tively rather than qualitatively.
2. Subsp. arachoides is the most variable of the three subspecies and comprises
three forms: the typical form (A) limited to the more arid parts of southern
Africa, namely, S.W.Africa, Botswana, the western Transvaal and the north-
western and northern Cape Province; the minute-haired form (B, C) repre-
sented in eastern southern Africa (the central and eastern Transvaal, Swaziland
and Natal as far south as the Tugela river valley, also in Rhodesia, and
Mocambique); the villous form (D) represented in the northern Transvaal (also
probably in Rhodesia).
It has not been considered necessary to cite specimens additional to those
listed by Brenan (1958).
It is interesting to postulate why subsp. micrantha should possess a stigma
different from that of both other subspecies and why its distribution should be
limited to India and central Africa and especially the latter where it breaks the
distributional continuity of the stylar type representative of subsp. italica and
arachoides. Can this be attributed to introgression by genetic material of a species
of Cassia present in India and Central Africa, but absent elsewhere from the range
of C. italica? C. tora in India and C. obtusifolia in Tropical Africa are closely
related to one another and have a stigmatic form that might account for that of
subsp. micrantha. All three taxa belong to the subgenus Senna, but to different
sections of this subgenus. All have been recorded as possessing races with 14
chromosomes in the gametophytic phase (Irwin & Turner, 1960). Experimental
breeding is required to test this postulate.
A duplicate of one of the syntypes of C. obovata Collad. var. pallidiflora
Dinter in Feddes Repert. 15: 355 (1918) (Dinter 366, Okahandja) is preserved at
the South African Museum Herbarium (SAM). This specimen agrees with other
specimens of subsp. micrantha from South West Africa cited by Brenan (1958)
and thus Dinter’s variety may now be accepted as falling within the limits of
subsp. micrantha. Brenan (l.c.) stated that no syntypes of Dinter’s variety were
still in existence at Berlin and duplicates had been looked for in vain in the
herbaria at Hamburg, the British Museum (Natural History) and Kew.
SCANNING ELECTRON MICROGRAPHS
The micrographs illustrating this paper were prepared in the following way:
representative stigmas and pieces of ovaries and mature leaves removed from
Genus Cassia in SA: Notes on Cassia italica 81
herbarium specimens were mounted on stubs using double-sided adhesive tape.
After drying for 24 hours in a desiccator, the specimens were coated in vacuum
with gold-palladium to a thickness of less than 150 A, then examined in a Hitachi
SSM2 scanning electron microscope at a voltage of 10 kV. The images were
observed at a magnification range 100- and photographed with a 35 mm camera.
KEY TO SUBSPECIES AND FORMS REPRESENTED IN SOUTHERN AFRICA
Stigma (usually visible even on developing fruits) expanded to exceed the width of the
style, asymmetrically trumpet-shaped (better developed adaxially than abaxially); style
3-4 mm long, curved but not + circinnate; flowers pale yellow to cream, brown
veined; racemes, when only basal flowers are open, 20-80 mm long, usually shorter
thankthetsubtendinpaleati cryin acres tess a) aatkersias nee Ses elolaanr subsp. micrantha
Stigma (usually visible even on developing fruits) hardly expanded (subterminally only
to slightly exceed style width) narrowing terminally to form an aperture usually
narrower than the style; style 6-7 mm long, + circinnate; flowers bright yellow, brown
veined only with age; racemes, when only basal flowers are open, mostly 60-150 mm
long, usually longer than the subtending leaf....................... subsp. arachoides
Ovarymolabroushir cee eis nate sets tis ac aueels atthe pics terse typical form (A)
Ovaryabearin SitriChOMeS rave soe cede ee ceHin eset a aucee ae ale ei saitasievayy lo Gleusllads, Sueimapels
Trichomes long, fine + straight, closely packed so as to form a villous
(DUIDCHCCNCCs s cd ba eonsboadonbouds eaes od negodoupooacep do villous form (D)
Trichomes short (microscopic) rather thick for their length, straight or curved,
closely packed when ovary very young ............ minute-haired form (B & C)
ACKNOWLEDGEMENTS
I wish to thank Miss L. Cowan, Mrs B. Loutit and Messrs V. Bandu, P. Evers,
B. Martin, R. Poonsamy and D. Tunnington for help in the preparation of
illustrations; Dr J. H. Ross for the checking of specimens, and the Curators of
Herbaria for the loan of material. Financial assistance from the Council of
Scientific and Industrial Research is gratefully acknowledged.
REFERENCES
BAKER, E. G., 1930. The Leguminosae of Tropical Africa. Ostend: Unitas Press, pp.
636-637.
BRENAN, J. P. M., 1958. New and noteworthy Cassias from Tropical Africa. Kew
Bull. 13: 239-243.
Burtt Davy, J., 1932. A manual of the flowering plants and ferns of the
Transvaal with Swaziland, South Africa 2: 325. London: Longmans, Green & Co.
Dinter, K., 1918. Index der aus Deutsch-Siidwestafrika bis zum Jahre 1917
bekannt gewordenen Pflanzenarten II. Feddes Rep. 15: 355.
Irwin, H. S. AND TURNER, B. L., 1960. Chromosomal relationships and taxono-
mic considerations in the genus Cassia. Am. J. Bot. 47: 309-318. ;
SCHREIBER, A., 1967. Caesalpiniaceae. In: Prodromus einer flora von Siidwest-
afrika 59: 11.
VERDCOURT, B., 1970. Studies in the Leguminosae—Papilionoideae for the Flora
of Tropical East Africa: III. Kew Bull. 24 (3): 379-447.
Jl S. Afr. Bot. 44 (1): 83-87 (1978)
NUCLEOLAR HETEROCHROMATIN IN ENCEPHALARTOS
D. J. MOGFORD
(Department of Plant Sciences, Rhodes University, Grahamstown)
ABSTRACT
Using a denaturation-reannealing-fluorescence technique to illustrate the position of
heterochromatin, it is shown that during interphase in the root tip cells of Encephalartos a
close association occurs between heterochromatin and the nucleoli. The possible signifi-
cance of this is discussed.
UITTREKSEL
NUKLEOLER HETEROCHROMATIEN IN ENCEPHALARTOS
Die gebruik van ’n denaturering-hergloei-fluoreserende tegniek om die posisie van
heterochromatien te illustreer, toon dat daar in die wortelpunt selle van Encephalartos
gedurende die interfase *n nou assosiasie tussen die heterochromatien en die kernliggaam-
pies is. Die moontlike betekenis hiervan word bespreek.
INTRODUCTION
The functions of heterochromatin, i.e. the late-replicating fraction of DNA
present as condensed “‘chromocentres’’ in interphase nuclei, remain obscure. This
is despite widespread application of the recently discovered fluorescence and
denaturation-reannealing techniques which indicate the location of heterochroma-
tin bands in metaphase chromosomes (Caspersson ef al., 1968; Pardue and Gall,
1970).
In general, heterochromatin continues to be associated with developmental
effects of the quantitative type. However, great diversity occurs not only in the
nature of its effects but also in the nature of heterochromatin itself.
For example, Vosa (1970) demonstrated that there exist in plants at least four
types of heterochromatin, characterised variously by enhanced or reduced
quinacrine fluorescence and by positive or negative sensitivity to cold. The various
responses of heterochromatin to denaturation-reannealing treatment followed by
Giemsa staining add further complexity to the situation, such that in Allium flavum
alone there exist four types of heterochromatin as characterised by their various
responses to fluorescence and Giemsa staining (Vosa, 1973).
Of late, it has proved possible to obtain new evidence on the functions of
heterochromatin by using the new techniques to identify the location of hete-
rochromatin in interphase nuclei. Thus Mogford (1977) demonstrated that hete-
rochromatin is involved in an end-to-end fusion of chromosomes in onion
Accepted for publication 14th July, 1977.
83
84 Journal of South African Botany
interphase nuclei, with the various consequences this implies for gene transcription
and meiotic pairing.
The present study was designed to investigate the relationship between
heterochromatin and nucleolar synthesis in the interphase nuclei of Encephalartos
species.
MATERIAL AND METHODS
Investigations were performed on growing root tips from seedling specimens
of Encephalartos lehemanni Ecklon ex Lehm. and E. caffer Miq.
Nucleoli were studied by means of phase-contrast illumination using visible
light, while heterochromatin was revealed using the combined denaturation-
reannealing-fluorescence technique described previously (Mogford, 1977). The
relationship between heterochromatin and the nucleoli was observed by means of
simultaneous reflected-light fluorescence and transmitted-light phase contrast,
using a Zeiss fluorescence microscope.
IGaale
Metaphase chromosomes of E. lehemanni, prepared as indicated in text. Arrows
indicate positions of heterochromatin; note the heterochromatic (and heteromorphic) satel-
lites of the nucleolar organising chromosomes, marked by the larger arrows. X1349.
Nucleolar Heterochromatin in Encephalartos
85
Fic. 2.
E. lehemanni, X1349.
Fic. 3.
7 E. caffer, X843.
Fics 2 & 3
Interphase nuclei of Encephalartos, showing relationship between
tin and nucleoli. Photographed using combined fluorescence and phase contrast as described
in text.
heterochroma-
86 Journal of South African Botany
RESULTS
Heterochromatin in E. lehemanni was found to be present on the nucleolar
chromosome pair, principally in the vicinity of the nucleolar organisers, and also
on certain of the other chromosomes (Fig. 1).
In both species, the heterochromatin visible during interphase was found to be
strongly localised at positions along the borders of the nucleoli (Figs 2 & 3). It
was assumed that this heterochromatin was that of the nucleolar chromosome pair,
though the possibility that other heterochromatin was involved as well could not
be ruled out since fusion of chromocentres during interphase is a common
phenomenon.
DISCUSSION
The occurrence of heterochromatin in the vicinity of the nucleolar organisers,
even in the quantities recorded in the present observations, is not by itself unusual.
What is unusual is the extremely conspicuous association of heterochromatin with
nucleoli in the interphase nuclei. This association in cycads may to some extent be
inferred by conventional staining (Marchant, 1968), but is demonstrated with
particular clarity by the present technique.
The relationship raises again the possible involvement of heterochromatin in
nucleolar synthesis, for the consistency of the association is greater than that
which might be expected simply from the location of heterochromatin in the
vicinity of the secondary constrictions.
Brown (1966) suggested that heterochromatin in the vicinity of the nucleolar
organiser might serve as an inert region of chromosome, separating off the intense
ribosomal RNA synthetic activity of the nucleolus organiser itself from the
activities of the remainder of the chromosome.
An alternative possibility is that heterochromatin itself might be involved in
nucleolar synthesis. Certainly, such an intense and specific role would be consis-
tent with the high degree of base repetition which modern studies indicate as
characterising heterochromatin. Such repetition, indeed, would be consistent with
the general quantitative action of heterochromatin. The difficulty here is that DNA
in the heterochromatic state is normally assumed to be non-synthetic, though as a
generalisation this is now less tenable in view of recent evidence for a differential
amplification of certain heterochromatic DNA in both plant and animal species
(Schweizer and Nagl, 1976). Of particular relevance is the observation of a
differential amplification of nucleolus-associated heterochromatin in both the
beetle Dytiscus marginalis and the cricket Acheta domestica (John and Lewis,
1975). The possibility of a synthetic role cannot therefore be excluded.
ACKNOWLEDGMENT
May I thank Mr Cameron of the Grahamstown Botanical Garden for the loan
of cycad seedlings.
Nucleolar Heterochromatin in Encephalartos 87
REFERENCES
Brown, S. W., 1966. Heterochromatin. Science, N.Y. 151: 417-425.
CASPERSSON, T., FARBER, S., FOLEY, G. E., KupyNowski, J., Mopest, E. J.,
SIMONSSON, E., WAGH, U. and ZECH, L., 1968. Chemical differentiation along
metaphase chromosomes. Expl. Cell Res. 49: 214-222.
Joun, B. and Lewis, K. R., 1975. Chromosome hierarchy. Oxford University
Press.
MarcHANT, C. J., 1968. Chromosome patterns and nuclear phenomena in_ the
cycad families Stangeriaceae and Zamiaceae. Chromosoma 24: 100-134.
Mocrorp, D. J., 1977. Chromosome associations in onion root tip nuclei. J/ S.
Afr. Bot. 43: 97-102.
PARDUE, M. L., and Gat, J. C., 1970. Chromosome localisation of mouse
satellite DNA. Science 168: 1356-1358.
SCHWEIZER, D., and NAGL, W., 1976. Heterochromatin diversity in Cymbidium,
and its relationship to differential DNA replication. Expl. Cell Res. 98: 411-423.
Vosa, C. G., 1970. Heterochromatin recognition with fluorochromes. Chromo-
soma 30: 366-372.
Vosa, C. G., 1973. The enhanced and reduced fluorescence bands and_ their
relationship to the Giemsa patterns in Allium flavum. Nobel Symposium 23:
156-158.
JI S. Afr. Bot. 44 (1): 89-91 (1978)
ee
TWO NEW SPECIES AND A NEW COMBINATION IN THE GENUS
KALANCHOE
H. R. TOLKEN
(Botanical Research Institute, Pretoria)
ABSTRACT
K. neglecta Toelken, K. rubinea Toelken and the new combination K. luciae Hamet
subsp. montana (Compt.) Toelken are published.
UITTREKSEL
TWEE NUWE SPESIES EN ’N NUWE KOMBINASIE IN DIE GENUS KALANCHOE
K. neglecta Toelken, K. rubinea Toelken en die nuwe kombinasie K. /uciae Hamet
subsp. montana (Compt.) Toelken word gepubliseer.
Raymont Hamet worked for many years on the taxonomy of the genus
Kalanchoe. He was, however, unable to complete a revised version of his early
monograph published in 1907, and a number of names used by him on determina-
vit labels were not validly published. The need thus arose to re-evaluate these taxa
and to validly publish them in order to avoid confusion.
Kalanchoe luciae Hamet subsp. luciae
K. luciae Hamet in Bull. Herb. Boissier ser. 2,8: 256 (1908); Hamet et Marnier-
Lapostolle, Gen. Kalanchoe au Jard. Bot. Les Cedres 91, figs 112-114 (1964).
Type: Transvaal, Spelonken, Junod s.n. (G, holo.).
Plants often up to 1,5 m high and glabrous. Leaves obovate to orbicular rarely
oblanceolate. Sepals 2,5-4-5) mm long, glabrous. Squamae usually square to
transversely oblong.
— subsp. montana (Compt.) Toelken, comb. nov.
K. montana Compt. in Jl S. Afr. Bot. 33: 295 (1967). Type: Swaziland, near
Devil’s Bridge, Compton 29 471 (NBG, holo.!).
Plants rarely more than | m high and usually hairy. Leaves narrowly oblanceo-
late to rarely obovate. Sepals 5—6,5(-7) mm long, with scattered hairs. Squamae
oblong. :
The subsp. montana is usually found associated with rock outcrops in sour
grassveld while the typical subspecies grows on rocky upper slopes of hills in
bushveld.
ee ee
Accepted for publication 28th September, 1977.
89
90 Journal of South African Botany
Kalanchoe neglecta Toelken, sp. nov. a K. rotundifolia Haw. thyrso corymboso
denso et foliis profunde cordatis; a K. hirta Harv. lobis angustis corollae et foliis
profunde cordatis differt.
Herbae perennes ramis erectis, per florescentia usque ad 1 m altis et 15 mm in
diametro, glabrae. Folia petiolata, glabra; petioli 20-50(-80) mm longi, leviter
sulcati supra; laminae ovatae 35-80(-130) mm longae, profunde cordatae vel
peltatae, integrae vel leviter sinuatae, patelliformes vel planae, virides. Inflores-
centia thyrsus corymbosus floribus multis in dichasiis densis. Calyx lobis anguste
lanceolatis, (1,5—)2—3 mm longus, glaber, viridis. Corolla glabra, tubo paene tereti
6-7,5 mm longo; lobi anguste lanceolati et aceres, 2,5—3,5 mm longi, vitellini vel
aurantiaci. Stamina verticillis duobus longitudine inaeqati, antheris flavis 0,5—0,6
mm longis et quoque appendicula terminali indistincta. Sqguamae lineares 1,6—2,5
x 0,2-0,3 mm, gradatim decrescentes ad apices, vix crassae, pallide flavae.
Carpella stylis brevibus tenuibus et stigmatibus terminalibus; ovarium ovulis
multis quoque c. 0,5 mm longis.
Type: Natal, Sordwana Bay, Vahrmeijer & Tolken 535 (PRE, holo.!).
Perennial herbs with erect branches up to 1 m high and 15 mm in diameter when
in flower, glabrous. Leaves petiolate, glabrous; petioles 20-50-80) mm long,
slightly grooved above; lamina ovate 35-80(-130) mm long, deeply cordate or
peltate, entire or slightly sinuate, flat to patelliform, green. Inflorescence a
corymbose thyrse with many flowers arranged in dense dichasia. Calyx with lobes
narrowly lanceolate, (1,5—)2-3 mm long, glabrous, green. Corolla glabrous, with
tube almost terete and 6-7,5 mm long; lobes narrowly lanceolate and sharply
pointed, 2,5-3,5 mm long, yellow to orange. Stamens in two whorls of unequal
length, with yellow anthers 0,5—0,6 mm long and each with an indistinct terminal
appendage. Squamae linear, 1,6-2,5 X 0,2-0,3 mm, gradually tapering towards
the apices, scarcely fleshy, pale yellow. Carpels with short thin styles and terminal
stigmata; ovary with many ovules each c. 0,5 mm long.
Growing in small groups but never common, on sandy soil in north-eastern
Natal.
Hamet used the name K. rotundifolia var. peltata on determinavit labels but
this name cannot be used at species level because of K. peltata Baill.
Kalanchoe rubinea Toelken, sp. nov. a K. longiflora Schltr. ex J. M. Wood
ramibus teretibus et foliis plicatis recurvatis rubinescentibus differt.
K. longiflora var. coccinea Marnier-Lapostolle ex Jacobsen, Handb. Succ. Pl. 2:
652, fig 866 (1960); Sukk. Lex. 253, t. 105,1 (1970), non rite publicatum.
Herbae perennes ramibus decumbentibus teretibus sed inflorescentiis erectis in
flore usque ad 1 m altis. Folia quoque plerumque petiolo, glabra; petiolus
(8—-)15-30(-45) mm longus, paene teres sed sulco non profundo supra; lamina late
Two new species and a new combination in the Genus Kalanchoe 91
elliptica vel ovata, 50—70(—100) mm longa, 40—60(-80) mm lata, cordata in foliis
infernis sed plus minusve cuneata in foliis supernis, longitudinaliter plicata et plus
minusve recurva, lobis quoque 1—3(4) dentibus obtusis, viridis vel atrorubinea.
Inflorescentia thyrsus corymbosis dichasiis multis. Calyx lobis triangulari-
lanceolatis, (1,5-)2-3(—5) mm longus, glaber, flavo-viridis. Corolla glabra, tubo
9-14-15) mm longo et quadrangulari; lobi late ovati vel paene orbiculares et
mucronibus terminalibus, 2,5-4 mm longi, minute papillosi, flavi. Stamina ver-
ticillis duobus longitudine inaequati, antheris flavis 0,7-1 mm longis et quoque
appendicula terminali gracili. Squamae lineari-lanceolatae, 1,6-4 x 0,8-1,1 mm,
apicibus gradatim constrictis, vix succulentae, pallide flavae. Carpella stylis
papillosis gracilibus et stigmatibus terminalibus prominentibus; ovarium 50-80
ovulis quoque 0,6—0,8 mm longis.
Type: Transvaal, Soutpansberg, Galpin 14 934 (PRE, holo.!).
Perennials with decumbent branches but erect inflorescenes up to 1 m high when
in flower, with terete branches. Leaves usually petiolate, glabrous; petiole (8-)
15-30(-45) mm long, almost terete but with indistinct groove above; lamina
broadly elliptic to ovate, 50-70(-100) mm long, 40-60(—-80) mm broad, cordate
on lower leaves, cuneate in upper leaves, folded length-wise and more or less
recurved, with lobes each with 1-3(-4) obtuse teeth, green to deep ruby red.
Inflorescence a corymbose thyrse with many dichasia. Calyx with lobes triangular-
lanceolate (1,5-)2-3(-5) mm long, glabrous, yellowish-green. Corolla glabrous,
with tube 9-13(-15) mm long and quadrangular; lobes broadly ovate to almost
orbicular and with terminal mucro, 2,5-4 mm long, minutely papillose, yellow.
Stamens in two whorls of unequal length, with broad yellow anthers 0,7—-1 mm
long and each with a slender terminal appendage. Squamae_linear-lanceolate,
1,6-4 x 0,8-1,1 mm, gradually tapering towards the apex, scarcely fleshy, pale
yellow. Carpels with slender papillose style and prominent terminal stigmas, ovary
with 50-80 ovules each 0,6—0,8 mm long.
K. rubinea occurs along the Eastern Transvaal escarpment from eastern
Swaziland to the Soutpansberg.
This species, which is widely used in horticulture, was always confused with
K. longiflora, an endemic species of central Natal.
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JI S. Afr. Bot. 44 (1): 93-96 (1978)
BooK REVIEWS
GLOSSARY OF GENETICS AND CYTOGENETICS, CLASSICAL AND MOLECULAR, by R.
Rieger, A. Michaelis and M. M. Green, with pp. 647. Fourth completely revised
edition. ISBN 3-540-07668-9. Berlin-Heidelberg-New York, Springer-Verlag,
1976. US $14.80.
The new Glossary is an impressively efficient looking book with some 3 500 entries and
1 700 references. The print is clear and readable and the text is liberally interspersed with
clear schematic line drawings. Arrows (—) indicate cross-references. Its size, 20 x 14 x 3
cm, makes it easy to handle and, although the review copy has a soft cloth cover, it is
strongly bound and should last well.
The third edition (the first English edition) has been in our department for many years
and I have consulted it frequently. However, on being asked to review the new edition I
realized that consulting and reviewing this kind of book are quite different. One usually
consults a glossary for the meaning of terms in fields of the subject with which one is not
familiar. On the other hand, in assessing the value of the book as reviewer, one is restricted
to the assessment of terms in one’s own specialized field. My impression of this book after
having looked at it as a reviewer in the field of quantitative genetics is one of disappoint-
ment with the quality and the accuracy of the text. I present the following examples in
support of this criticism.
p. 24 allozygote (McKusick, 1973)—‘‘a genetic compound, i.e., heterozygous for different
alleles’’. This definition is incomplete and misleading. Crow and Kimura (1970)
follow the notation of Cotterman in designating an individual whose two homolog-
ous genes are identical by descent as autozygous. If the two alleles are of
independent origin, the individual is said to be allozygous. Allozygotes can therefore
be either homozygotes or heterozygotes. The term autozygote is not included in the
Glossary. This is a pity because it could have been used to avoid the muddle in
describing the effects of inbreeding on page 290. In passing, McKusick (1973) is not
included in the bibliography.
p. 53 breeding value. In terms of the definition given here it is incorrect to omit the factor
of 2.
p. 111 coancestry. Falconer (1960) has been misquoted. The definition should refer to the
degree of relationship by descent, not to the degree of identity by descent of the two
parents. Also it is incorrect to say that coancestry may be used in place of the
coefficient of relationship.
p. 112 coefficients of coancestry, kinship and parentage are given on the same page. They
refer respectively to individuals I & L, A & B, and X & Y and different words are
used to describe these synonyms. It is an apt example of what Crow and Kimura
(1970) have described as a ‘“‘bewildering plethora of alternative names’ .
_ 112 coefficient of relationship—‘‘the probability that two individuals have inherited a
’ oe ae a panGGn cane or the proportion of all their allelic genes that
have been inherited from common ancestors’’. This merely adds to the general
bewilderment and the situation is certainly not clarified by the definition of genetic
relationship given on page 243.
93
94 Journal of South African Botany
p. 124 correlated response (Wigan and Mather, 1942)—‘‘a change in one character
(phenotype) occurring as an incidental consequence of selection for a seemingly
independent character’. This represents one of two major schools of thought. In
this ‘‘linkage school’’ the correlated response has low predictability. The “‘pleio-
tropy school’’ uses genetic correlation to predict correlated response. Omitting the
latter is inexcusable irrespective of one’s own affiliation. It is interesting to note
that genetic correlation is not listed in its own right but is included with character
(p. 74) and phenotype (p. 417). In each case both pleiotropy and linkage are given
as possible causes of genetic correlation. It is interesting to note also that the
following statement is made on page 494 under the listing of selection:
‘Artificial selection applied to one character almost always leads to changes in
others (‘‘correlated response’’)’’. Again, in passing, Wigan and Mather (1942) is
not included in the bibliography.
p. 252 genetic value—‘‘the phenotypic value an individual would have if there were no
environmental variation and no dominance effects’’. This is a very strange defi-
nition. It is cross-referenced to genetic variance (p. 254) where it is said that,
“Genetic variance can be divided into two components: 1. g.v. resulting from
differences between homozygotes (additive g.v.); 2. g.v. resulting from specific
effects of various alleles in heterozygotes (dominance variance)’’ and that, ‘“Dom-
inance here refers to deviations of the heterozygote from the two homozygotes’’.
When presented with this kind of draft by a student I find myself suggesting gently
but firmly that he re-consult his references, that he give more thought to the matter
and that he rewrite the section concerned.
p. 256 genotype-environment interaction. Although no reference is given it is fairly clear
that Falconer’s (1960) provisional clearance of the decks of complications when
discussing the additivity of variance components has been rather ineptly summa-
rized. No further attempt has apparently been made by the compilers to understand
the concept. |
p. 267 heritability. It is stated inter alia that, ‘‘Heritability is a measure of the amount of
genetic variability, excluding that expressed by heterozygotes. . .”’. This muddle |
arises I think from the assumption, evidenced in the Glossary in the definitions of
genetic value and genetic variance, that pure additivity of gene action is a
prerequisite for additive genetic variance.
p. 290 inbreeding coefficient (Wright, 1929). It is stated that, ‘‘If the frequencies of two
alleles (A and a) in a population are pA and qa and the inbreeding coefficient is F,
then the chance that an individual will be genotypically AA is pF. . .. The chance
that the two alleles at that locus will not be identical by descent is 1—F’’. This is
incorrect. The expected frequency of AA autozygotes is pF while the expected
frequency of AA allozygotes is p? (I-F). In passing, the reference to Wright (1929)
is inappropriate and the page number given in the bibliography is incorrect.
p. 339 maternal effect—‘any nonlasting effect of the maternal genotype or phenotype on
the immediate offspring.’’ I do not think a maternal effect is necessarily of limited
duration nor do I think that a definition of maternal effect without mentioning
maternal environment is adequate. I think ‘‘immediate’’ is an inappropriate choice
of word in so far that it is used with a specific meaning by both Penrose (1954) and
Robson (1955) in describing maternal effects.
p. 44
Nn
progeny selection and progeny test. The definitions are not precise and there is no
mention of or cross-reference to breeding value. The ingredients for predicting
selection response are given without saying why they should be considered.
————————— a
Book Reviews 95
p. 496 selection response. After nearly three pages devoted to various forms of natural and
artificial selection including definitions of selection coefficients, pressures, differen-
tials and limits, selection response reads, in full: ‘‘the gain in liability due to
selection which is a function of the heritability of liability, assuming that genotype-
environment interactions are absent. If A is the mean liability of affected individuals
(those in the tail of the general population distribution), and R the mean of their
offspring, then the realized gain in liability is R-A’’. This to say the least is
inappropriate since no prior reference or cross-reference is made to threshold
characters. A few words concerning the predictability of selection response could
surely have been included?
In fairness to the compilers I feel I must repeat that I am not competent to judge the text
as a whole. At the same time I do not think they are competent in the field of quantitative
genetics and it is a pity they did not include in their team one who is.
J. S. ALLAN
KRYPTOGAMEN. BLAUALGEN, ALGEN, PILZE, FLECHTEN, by K. Esser, with pp.
573 + xvi, illus. Berlin-Heidelberg-New York, Springer-Verlag, 1976. DM58.00
Even though this book is bound in a soft cover it is a sturdy volume of convenient size
(16,5 x 24 cm). The text appears to have been typed and reproduced photographically on
good quality white glossy paper and is clear and easy to read. Two type sizes have been
employed, the smaller one instead of italics, without creating an unbalanced effect.
Except for occasional spelling errors there is little fault to be found with the accuracy of
the text. It is, however, also important to remember that taxonomic systems and arrange-
ments are subject to personal interpretation, and a reader may be surprised at finding some
taxa included in unexpected categories. Some taxonomic names which have been rejected
for somé time are again used in this book. By and large, however, the taxonomic
framework followed is acceptably up to date.
There is an abundance of illustrative material. The drawings are well done and clear, as
are the diagrams. The photographs are on the whole well chosen but are unfortunately
rather soft, with few definite blacks or whites. The overall impression is thus of rather dull
illustrations which lack the brilliance they could have had. A number of photographs have
also been retouched to accentuate desired areas. This could mislead a student into believing
that the detail he is looking for on a microscope preparation is more obvious than it in
actual fact is.
There are comparatively few references in the text. Those that are used are given as
footnotes and not combined into a single list. This does mean that to find a reference which
has previously been quoted can prove extremely troublesome. There are, however, compre-
hensive lists of suggested references, grouped under a variety of headings, at the back of the
book. These are extremely useful to student readers, but do not compensate for the
frustrations referred to above. ; ne
The index is comprehensive and detailed and there is also a list of taxa, an essential in a
book of this nature. fj
The book is aimed primarily at teachers of cryptogamic botany, would be very useful
for postgraduate students at Honours level but is not suitable for the unaided undergraduate.
The research worker. will not find much in it. Certain sections would not apply to South
African conditions but on the whole the information given could be applicable. However, as
the book is written in the German language, its local appeal is likely to be extremely
limited. Despite its attractions the text does not offer so much that the non-German
speaking reader will find it worth his while to translate it. a ; “of
This book is not a textbook in the normally accepted sense. It is intended to give a brie
review of the basic facts about the blue-green algae, the algae, the fungi and the eeu
followed by a section on technology where fixatives, media, preservatives, sampling an
96 Journal of South African Botany
handling techniques, etc. are given. The bulk of the volume (approximately 480 pages) is
then devoted to detailed discussions of individual organisms representative of the various
taxonomic subdivisions.
Techniques for finding and isolating organisms are followed by life cycle diagrams,
recommendations on cultivation and examination and, in most cases, photographs or
drawings of the various stages. Suggestions as to possible genetic experiments or other
demonstrations are also plentiful.
Whereas the book is thus extremely useful to the lecturer planning a course in
cryptogamic botany, the average student will find it difficult to obtain a coherent picture of
the organismic groups covered without extensive additional reading or assistance. As the
book is written for European conditions the South African lecturer will find some of the
suggested experimental organisms or programmes inappropriate.
There is, furthermore, a very important gap in the treatment of the fungi, and that is a
totally inadequate discussion of the Fungi Imperfecti. They are a large and successful group
of fungi and there are dynamic developments which are not even mentioned. Only eight
pages are devoted to the imperfect fungi and more than three of these to the dermatophytes.
The book ends with a useful section, admittedly less so for South African readers,
containing addresses of culture collections and suppliers of teaching films.
Despite its shortcomings, a highly interesting book.
K. T. VAN WARMELO
———
JI S. Afr. Bot. 44 (2): 97-101 (1978)
THE REPRODUCTIVE POTENTIAL OF ERICA JUNONIA BOLUS:
MEGASPOROGENESIS AND MEGAGAMETOGENESIS
B. L. ROBERTSON and JUDITH E. McNAUGHTON
(Department of Botany, University of Port Elizabeth)
ABSTRACT
The embryological study of Erica junonia forms part of an investigation into the
inadequate propagation of the species. The ovule is anatropous, unitegmic and tenuinucel-
late. The archesporial cell functions directly as the megaspore mother cell. Embryo-sac
development is monosporic and conforms to the Polygonum-type of the Supra Homotypic
Category. Starch grains are present in the integument and an endothelium lines the chalazal
region. Observations thus far suggest no restrictive effect on the reproduction of E. junonia.
UITTREKSEL
DIE VOORTPLANTINGSPOTENSIAAL VAN ERICA JUNONIA BOLUS: MEGASPO-
ROGENESE EN MEGAGAMETOGENESE
Die embriologiese studie van Erica junonia maak ’n deel uit van ’n ondersoek na die
gebrekkige voortplanting van die spesies. Die saadknop is anatroop, tenuinusellér en besit
‘n enkele integument. Die argesporiale sel funksioneer direk as die megaspoormoedersel.
Embriosakontwikkeling is monospories en geskied volgens die Polygonum tipe van die
Supra-Homotipiese Kategorie. Styselkorrels is teenwoordig in die selle van die integument
en die chalazagedeelte van die embriosak word deur ’n endotelium omhul. Hierdie studie
van megasporogenese en megagametogenese toon geen beperkings op die voortplanting van
E. junonia.
INTRODUCTION
The large-flowered variety of Erica junonia, reputed to be the most magni-
ficent of the ericas, occurs only on a single mountain peak in the Cold Bokkeveld
north of Ceres (Baker & Oliver, 1967). It flowers from November to January.
Germination of seeds has been unsuccessful, but the species can be propagated by
means of cuttings. However, these vegetatively propagated plants do not set seed.
The inadequate propagation of E. junonia led to its being declared an
endangered species by the C.S.I.R.’s Working Group on Rare and Endangered
plant species. This study forms part of an attempt to pinpoint the locus which
blocks the pathway to successful propagation.
MATERIAL AND METHODS
Flowers of E. junonia were collected from their natural habitat near Ceres and
fixed in formalin-acetic acid-alcohol. Dehydration was carried out in an ethyl
alcohol/tertiary butyl alcohol series and the material was embedded in Carowax
Accepted for publication 19th July, 1977.
97
98 Journal of South African Botany
(melting point 55 °C). Ovaries were sectioned on a rotary microtome at 10 wm and
stained with safranin and fast green (Sass, 1958).
RESULTS AND DISCUSSION
Megasporogenesis
Numerous ovules are contained in each of the four locules. Their placentation
is axile. Each ovule is anatropous (Fig. 2) and has a single, large integument (Figs
1 & 2). According to Davis (1966) these characteristics are reasonably constant for
the Ericaceae. When the embryo sac reaches maturity the epidermis of the
integument becomes completely tanniniferous.
———
~~
Megasporogensis in E. junonia
Fic. 1
Young ovule showing archesporial cell
(a = archesporial cell, i = integument) (< 600)
Fic. 2
ue Megaspore mother cell. Note the absence of nucellus
(i = integument, me = megaspore mother cell, t = tanniniferous cells) (<380)
Fic. 3
Heterotypic division of meiosis (x 380)
Fic. 4
Tetrad of megaspores in which three micropylar ones are degenerating
(functional megaspore arrowed) (x 380)
The reproductive potential of Erica junonia 99
At a very early stage in ovule development a single hypodermal cell becomes
prominent. This is the archesporial cell (Fig. 1) and it functions directly as the
megaspore mother cell. There are no parietal layers between the nucellar epi-
dermis and the megaspore mother cell and so the ovule can be classified as
tenuinucellate. According to Maheshwari (1950) there are two types of tenuinucel-
late ovules, namely, those in which the integument primordia are situated near the
apex of the nucellus and a second type in which the integument originates at the
base of the nucellus. In E. junonia the latter type occurs.
The nucellus is only three cell layers wide and is ephemeral. Its degeneration
occurs rapidly during the development of the megaspore mother cell. This cell
enlarges (Fig. 2) at the expense of the nucellus so that at maturity it is bordered
completely by the integument only.
Fig. 3 shows the metaphase stage of the heterotypic division during meiosis of
the megaspore mother cell. After completion of the homotypic division, a linear
tetrad of megaspores is formed (Fig. 4).
Megagametogenesis
Of these four megaspores only the chalazal one persists (Fig. 5) and functions
as the embryo-sac initial. The development of the embryo-sac is therefore
monosporic. The nucleus of the functional megaspore is centrally situated and
until this stage the embryo-sac exhibits no polarity.
Division of this nucleus results in the formation of a two-nucleate embryo-sac
(Fig. 6). One of these nuclei, the egg-apparatus mother nucleus, migrates towards
the micropyle while the other nucleus, the primary antipodal nucleus, moves to the
chalazal end. Because polarity of the embryo-sac is established at this stage of
development, the embryo-sac of E. junonia conforms to the Supra Homotypic
Category (Swamy & Krishnamurthy, 1975).
Once polarity has been achieved, the embryo-sac enters the elaboration phase.
Each of the two nuclei divides, producing a four-nucleate embryo-sac (Fig. 7) and
following yet another division, the embryo-sac contains eight nuclei.
These eight nuclei now undergo organization into the constituents of a mature
embryo-sac. At the micropylar end the tripartite egg apparatus consists of an egg
cell flanked by two synergids which soon become densely cytoplasmic (Fig. 8).
The micropylar region of the embryo-sac becomes spherical and enlarges at the
expense of the integumentary tissue, the cells of which contain numerous starch
grains. Davis (1966) reports the occurrence of starch grains in the embryo-sacs of
many ericaceous species.
A well-differentiated endothelium lines the chalazal region of the embryo-sac
(Figs 9 & 10)—the function of this endothelium is not clear. Opposite the point
where the endothelium layer terminates the secondary embryo-sac nucleus occurs
(Fig. 9).
Three antipodal cells are found in the chalazal region of the embryo-sac. Each
100 Journal of South African Botany
Megagametogenesis in E. junonia
Fic. 5
Uni-nucleate embryo-sac (d = degenerating megaspores) (380)
FIG. 6
Two-nucleate embryo-sac
(em = egg apparatus mother cell, pa = primary antipodal nucleus) (< 380)
Fic. 7
Elaboration phase showing four-nucleate embryo-sac (X 380)
Fic. 8
Egg apparatus (e = egg cell, synergids arrowed) (<380)
Fic. 9
Mature embryo-sac
(di = degenerating integument cells, containing starch grains, en = endothelium,
Ss = synergid, sn = secondary embryo-sac nucleus) (380)
Fic. 10
Mature embryo-sac (a = antipodal cells, sn = secondary embryo-sac nucleus) (380)
The reproductive potential of Erica junonia 101
cell has a definite cell wall (Fig. 10). The antipodals are derived from a single
primary antipodal nucleus and therefore the embryo-sac of E. junonia is of the
Polygonum type (Swamy & Krishnamurthy, 1975).
This study shows that the processes involved in the attainment of haploidy, the
establishment of polarity and the maturation of the embryo-sac appear to take
place normally. Up to this stage, therefore, there is no restriction on the
propagation potential of E. junonia.
ACKNOWLEDGEMENTS
The authors wish to express their gratitude to Mr J. Winter who was
responsible for the collection of plant material.
REFERENCES
Baker, H. A. and OLiver, E. G. H., 1967. Ericas in Southern Africa. Cape Town: Purnell
& Sons.
Davis, Gwenda L., 1966. Systematic embryology of the Angiosperms. New York:
John Wiley & Sons, Inc.
MaAHEsHwakrl, P., 1950. An introduction to the embryology of Angiosperms. New
York: McGraw-Hill Book Company, Inc.
Sass, J. E., 1958. Botanical microtechnique. Iowa: Iowa State University Press.
Swamy, B. G. L. and KrisHNAMURTHY, K. V., 1975. Embryo-sac ontogenies in
Angiosperms—an elucidation. Phytomorphology 25: 12-18.
he
i
JIS. Afr. Bot. 44 (2): 103-110 (1978)
EFFECT OF VARIOUS TREATMENTS ON GERMINATION OF
DORMANT SEEDS OF STRELITZIA REGINAE AIT.
H. A. VAN DE VENTER
(Department of Botany, University of Port Elizabeth)
ABSTRACT
The effects of a wide variety of dormancy-breaking treatments on the germination of S.
reginae seeds were investigated. Germination was improved after treating the seeds with
thiourea, various mercaptans, ascorbic acid and ethrel; by incubation in elevated oxygen
concentrations; and by removal of the tissues overlying the embryo in the seed. Treatments
which did not improve germination included incubation in elevated carbon dioxide
concentrations, leaching with water, various storage conditions, and treatment with urea,
indoleacetic acid, gibberellic acid, kinetin, various oxidizing agents, and respiratory
inhibitors. Low temperatures had a decidedly harmful effect on both dry and imbibed seeds.
UITTREKSEL
INVLOED VAN VERSKILLENDE BEHANDELINGS OP DIE ONTKIEMING VAN
STRELITZIA REGINAE AIT. SADE WAT IN ’N RUSTOESTAND VERKEER
Die invloed van ’n verskeidenheid rusbeéindigende behandelings op die ontkieming
van S. reginae sade is ondersoek. Ontkieming is verbeter nadat sade behandel is met
tio-ureum, verskeie merkaptane, askorbiensuur en ethrel; deur inkubasie in verhoogde
suurstofkonsentrasies sowel as na verwydering van die weefsels wat die embrio in die saad
oordek. Behandelings wat ontkieming nie verbeter het nie is: inkubasie in verhoogde
koolstofdioksiedkonsentrasies, loging van die sade met water, verskeie opbergingstoe-
stande, asook saadbehandeling met ureum, indoolasynsuur, gibberelliensuur, kinetien, ver-
skeie oksideermiddels en respiratoriese inhibeerders. Lae temperature het ’n besliste nadelige
uitwerking gehad op beide droé en geimbibeerde sade.
INTRODUCTION
Seeds of the acaulescent species of Strelitzia exhibit a condition of innate,
relative dormancy (van de Venter & Small, 1974). Evidence has been presented
that an inhibitor may be involved in this phenomenon (van de Venter & Small,
1975) but this does not preclude the possibility of contributions by other factors
since dormancy can be the result of a combination of factors (Villiers, 1972).
In an attempt to obtain more information on the mechanisms controlling
dormancy of S. reginae seeds, the preliminary approach of determining the effect
of various well-known dormancy-breaking treatments on these seeds was used. A
treatment which will promote the germination of dormant seeds can be expected to
have some bearing on the operative mechanism and it was envisaged that these
experiments would provide guidelines for further research.
Accepted for publication 7th September, 1977.
103
104 Journal of South African Botany
MATERIAL AND METHODS
Seeds of S. reginae, harvested from various plants in the East London area,
were used. The seeds were pooled and stored under ambient conditions in the
laboratory (ca. 20 °C). All experiments were conducted on seeds that were less
than a year old.
The seeds were sterilized and germinated in petri dishes as previously
described (van de Venter & Small, 1974). Due to limited quantity, only 25 seeds
per dish represented one replicate. The number of replicates varied between four
and eight, depending on the particular experiment.
Unless otherwise mentioned, only scarified seeds were used. Incubation
proceeded in the dark at 25 + 0,5 °C and counts were made after 14 days. The
seeds were scarified with emery paper to expose a few square millimeters of
endosperm.
Treatment of seeds with various chemicals was accomplished by immersing
them (after surface-sterilization) for 48 hours in sterile solutions of these com-
pounds, prior to incubation in petri dishes.
In experiments involving treatments with oxygen and carbon dioxide, seeds
were incubated in sterile, 130 mf-capacity conical flasks containing 4 mf sterile,
distilled water. Oxygen and carbon dioxide concentrations were adjusted to
required values by injecting specific volumes of these gases through the vaccine
caps sealing the flasks. Where an atmosphere of pure oxygen was required, flasks
were flushed with this gas for three minutes.
Arcsin transformation of germination percentages was carried out before
Statistical tests were applied and lowest significant difference (LSD) values
calculated according to the w-procedure of Tukey (Steel & Torrie, 1960).
RESULTS
Effect of light and scarification
Scarified and unscarified seeds were subjected to light and dark treatments in a
2 x 2 factorial design. Light was supplied from cool white fluorescent tubes at an
intensity of ca. 12 000 lux. Petri dishes of the dark treatment were wrapped in a
single layer of aluminium foil.
The results of this experiment (Table 1) show that scarification did not
improve germination to a significant extent. However, this treatment improved
water absorption by the seeds (Fig. 1). Light had a harmful effect on the
germination of unscarified seeds.
Effect of thiourea, urea, various mercaptans and ascorbate
Seeds were soaked in different concentrations of thiourea, urea, reduced
glutathione, L(+) cysteine, 2,3-dimercapto-1-propanol (BAL) and L(+) ascorbic
acid. All of these compounds, with the exception of urea, promoted germination at
certain concentrations (see Table 2).
aE a
Effect of various treatments on germination of dormant seeds 105
TABLE |
Germination percentage of scarified and unscarified seeds of S. reginae in light and dark
(Arcsin transformed values in brackets)
Light Dark Mean
Scarified seeds 31 (33,49) 25 (29,21) 28 (31,35)
Unscarified seeds 13 (20,20) 30 (31,96) 21 (26,08)
Mean 22 (26,85)
27 (30,58)
Effects: F Values: LSD
Light 1,85 —
Scarification 3,68 —
Lightx Scarification 8,50%* (10,55)
Effect of growth substances
Seeds were soaked in concentration series of gibberellic acid (GA;; 0,1—1 000
ppm), indoleacetic acid (1-1 000 ppm), kinetin (3-48 ppm) and ethrel (1 000—
5 000 ppm active ingredient). With the exception of ethrel, no significant
promotion of germination was found. The optimal concentration of ethrel was
seeds abraded
seeds not
abraded
Water absorption as % of original mass
0 24 48 72 ze sz0
Time (hours)
Fic. 1 ;
Effect of abrasion on water absorption by S. reginae seeds.
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Effect of various treatments on germination of dormant seeds 107
found to be 2 000 ppm. At this concentration 65% of the seeds germinated as
compared to 27 % in the control.
Effect of carbon dioxide and oxygen
The treatments applied in these experiments are listed in Table 3 and are
expressed in terms of volumes of CO, and Os in the flasks as a percentage of the
total gas volume.
TABLE 3
Effect of different carbon dioxide and oxygen concentrations on germination of S. reginae
seeds (Arcsin transformed values in brackets)
Carbon dioxide
Concentration Germination Concentration Germination
% % % %
0 (KOH in centre well) 3 20,0 (air) 2 (4,66)
24,1 2 (4,66)
0,03 (air) 27,9 3 (Gio)
Des 34,7 10 (18,09)
48 100,0 42 (40,24)
F value: 29,74**
LSD: (11,48)
The results of these experiments should be interpreted with caution as the
seeds in the different treatments were exposed to different gas pressures. Respira-
tory activity of the seeds could also have changed the original gas composition.
In spite of these technical restrictions it seems valid to conclude from the
results in Table 3 that whereas elevated CO, concentrations elicited no response,
elevated O, concentrations had a stimulatory effect on germination. Poor germina-
tion in the control and CO, treatments was probably the result of limiting oxygen
in the sealed flasks.
The beneficial effect of oxygen was confirmed in a subsequent experiment
using larger flasks (600 m£ capacity). Incubation in 100 % oxygen resulted in 64 %
of the seeds germinating within one week. Seeds in control flasks that were not
sealed, but stoppered with cotton-wool, showed a germination of only 11%.
Mechanical restriction of embryo growth
Scarified seeds were allowed to imbibe water for two days after which the
tissues covering the embryo in the proximal region of the seed were excised. As
sterile conditions were difficult to maintain during this so-called chipping oper-
ation, the seeds were not sterilized. The seeds were incubated in quartz sand kept
moist with daily applications of tap water. After three weeks, only eight per cent
108 Journal of South African Botany
of the seeds in the control treatment had germinated in comparison with the 53 %
germination attained by chipped seeds.
Effect of different storage conditions
Air-dry seeds were kept in cotton-wool stoppered flasks for four weeks at
room temperature (ca. 20 °C), 40 °C, 50 °C as well as in sealed 600 mf? flasks (at
ca. 20 °C) flushed weekly with pure oxygen. A stratification treatment (imbibed
seeds kept at 4 °C for four weeks) was also included.
None of the treatments resulted in an increased germination. Both storage at
50 °C and stratification proved to be decidedly deleterious, depressing germination
by 21 and 34 %, respectively, relative to the room-temperature control.
A subsequent observation on air-dry seeds that had been stored for 10 months
at 4 °C, showed that this treatment decreased germination by 18 % in comparison to
laboratory-stored seeds.
Effect of oxidizing agents and respiratory inhibitors
Treatments consisting of concentration series of KNO, (1-1 000 ppm), KNO,
(1-10 000 ppm), H,O, (2-8M with different exposure times), methylene blue
(10-+10 °M) and KCN (10~*-10 °M) failed to show any stimulatory effect on
germination. Germination was improved by a concentration of 10 *M sodium
azide, but the effect was not statistically significant.
Effect of leaching
Although there is evidence for the presence of a water-soluble inhibitor in the
seeds of S. reginae (van de Venter & Small, 1975), leaching of scarified seeds in
running tap water for six days failed to improve germination.
DISCUSSION
Judging from the variety of treatments which have been found to improve
germination of S. reginae seeds, their dormant condition seems to be a complex
phenomenon with possibly more than a single causative factor. It would be
premature to speculate on possible control mechanisms at this stage but, neverthe-
less, the results have implicated the involvement of various factors which might be
investigated more fully.
Thiourea, a well-known dormancy-breaking agent (Mayer & Poljakoff-
Mayber, 1963) was found to improve germination of S. reginae seeds. The fact
that various mercaptans, in contrast to urea, were able to improve germination,
suggests that the stimulatory effect of thiourea resulted from the tendency of this
compound to form a thio-imido tautomer which occurs in equilibrium with the
normal form of thiourea (Toole, Hendricks, Borthwick & Toole, 1956; Garrard &
Biggs, 1966). That the reductive capacity of these compounds is involved, was
suggested by the positive results with ascorbate.
Effect of various treatments on germination of dormant seeds 109
Paradoxically, the promotive effects of increased oxygen concentrations on
germination indicate the necessity of an oxidation reaction for the termination of
dormancy. Alternatively this result could simply be indicative of a permeability
barrier in the seed coat. The magnitude of the response to oxygen points to a
major involvement of this gas in the termination of dormancy and indicates that
this factor should be further investigated.
Roberts (1969, 1973) proposed that, in the seeds of certain species,
cytochrome oxidase competes for available oxygen with a dormancy-breaking
reaction. He postulated that operation of the pentose phosphate pathway is a
prerequisite for the termination of dormancy. The seeds of S. reginae, although
responding positively to oxygen and thiourea, failed to do so to electron acceptors,
respiratory inhibitors, storage in elevated oxygen tension, and stratification. These
results suggest that the mechanism, as proposed by Roberts, would not serve as a
hypothetical model for dormancy in Strelitzia seeds.
The view that dormancy is essentially under hormonal control, that is,
inhibitors interacting with growth promoters, has been expounded by authors such
as Mayer (1960) and Amen (1968). Control of this nature in the case of Strelitzia
is a distinct possibility and merits further attention. Evidence for the presence of
an inhibitor has been presented (van de Venter & Small, 1975) and the involve-
ment of ethylene in the termination of dormancy (as implied by the effect of
ethrel) is indicated by these results.
Possible correlations between the factors already mentioned and the mechani-
cal restriction of embryo growth by the seed coat are not clear at this stage.
Mechanical restriction by the seed coat is known to be involved in the seed
dormancy of various species (Randolph & Cox, 1943; Chen & Thimann, 1964,
1966; Chen, 1968).
The significance of the light x scarification interaction is also vague. Mayer &
Poljakoff-Mayber (1963) have reported that seed-coat damage can alter the
response of seeds to light.
ACKNOWLEDGEMENTS
Financial assistance from the Council for Scientific and Industrial Research
and the University of Port Elizabeth is acknowledged.
REFERENCES
Amen, R. D., 1968. A model of seed dormancy. Bor. Rev. 34: 1-31.
CHEN, S. S. C., 1968. Germination of light-inhibited seeds of Nemophila insignis.
Am. J. Bot. 55: 1177-1183.
CuHen, S. S. C. and Tuimann, K. V., 1964. Studies on the germination of
light-inhibited seeds of Phacelia tanacetifolia. Israel Jnl Bot. 13: 57-73.
CHEN, S. S. C. and THIMANN, K. V., 1966. Nature of seed dormancy in Phacelia
tanacetifolia. Science, N.Y. 153: 1537- 1538.
GARRARD, L. A. and Biccs, R. H., 1966. Thioamide-induced germination of
seeds of Prunus persica. Phytochem. 5: 103-110.
110 Journal of South African Botany
Mayer, A. M., 1960. Germination research at the Hebrew University Jerusalem,
Israel—A review. Indian J. Pl. Physiol. 3: 13-23.
Mayer, A. M. and PoLsJAKOrFF-MAYBER, A., 1963. The germination of seeds.
London: Pergamon Press.
RaNpDOoLpH, L. F. and Cox, L. G., 1943. Factors influencing the germination of
Tris seed and the relation of inhibiting substances to embryo dormancy. Proc. Am.
Soc. hort. Sci. 43: 284-300.
Roperts, E. H., 1969. Seed dormancy and oxidation processes. In: Dormancy
and survival: 161-192. Cambridge: University Press.
Roperts, E. H., 1973. Oxidative processes and the control of seed germination.
In: W. Heydecker (ed.), Seed ecology. Chap. 2. London: Butterworths.
SreeEL, R. G. D. and Torrie, J. H., 1960. Principles and procedures of statistics.
Ist ed. New York: McGraw-Hill Book Co.
TooLe, E. H., HeENpricKs, S. B., BortTHWwick, H. A. and Toore, V. K., 1956.
Physiology of seed germination. A. Rev. Pl. Physiol. 7: 299-324.
VAN DE VENTER, H. A. and SMALL, J. G. C., 1974. Dormancy in seeds of
Strelitzia Ait. S. Afr. J. Sci. 70: 216-217.
VAN DE VENTER, H. A. and SMALL, J. G. C., 1975. Evidence for the presence of
a germination inhibitor in seeds of Strelitzia Ait. JI S. Afr. Bot. 41(4): 211-223
Vittiers, T. A., 1972. Seed dormancy. In: T. T. Kozlowski (ed.), Seed biology.
Vol. Il. Chap. 3. New York: Academic Press.
PL ssn
JIS. Afr. Bot. 44 (2): 111-117 (1978)
CENTROMERIC HETEROCHROMATIN IN LACHENALIA TRICOLOR
(L.) THUNB.
D. J. MOGFORD
(Department of Plant Sciences, Rhodes University, Grahamstown)
ABSTRACT
The South African Lachenalia tricolor is notable for possessing large amounts of
heterochromatin. The heterochromatin is centromeric in position, and may be revealed
either by direct quinacrine staining or by denaturation-reannealing treatments. The signi-
ficance of this is discussed relative to the general occurrence and adaptive significance of
this type of chromatin.
UITTREKSEL
SENTROMERIESE HETEROCHROMATIEN IN LACHENALIA TRICOLOR (L.)
THUNB.
Die Suid-Afrikaanse Lachenalia tricolor word deur groot hoeveelhede heterochromatien
gekenmerk. Die heterochromatien is sentromeries in plasing en kan deur direkte beitsing
met quinakrien of met denaturasie-hervoeging behandeling aangedui word. Die betekenis
hiervan word bespreek met betrekking tot die algemene voorkoms en aanpasbaarheidsbete-
kenis van dié soort chromatien.
INTRODUCTION
Heterochromatin, i.e. the late-replicating fraction of DNA present as con-
densed ‘‘chromocentres’’ in resting nuclei, has been the subject of renewed
interest over recent years. This has followed the discovery that the positions of
heterochromatin bands in metaphase chromosomes may be revealed by their
differential fluorescence in ultraviolet light following treatment with certain
fluorochromes (Caspersson ef al., 1968), and by their differential staining with
Giemsa or orcein following denaturation-reannealing treatments (Pardue and Gall,
1970; Vosa, 1973a). The latter method derives from the highly repetitive base
constitution of heterochromatin, which results in its reannealing more completely
than the remainder. These techniques are of much wider applicability than the
older method based on the understaining of heterochromatin in chromosomes
exposed to low temperature (Darlington and La Cour, 1940), for only certain types
of heterochromatin show such cold sensitivity. ‘el
The new techniques have greatly increased the number of plant species in
which heterochromatin is known to occur. Heterochromatin has been demon-
strated, for example, in Vicia (Vosa and Marchi, 1972), Hepatica, Anemone
(Marks and Schweizer, 1974), and Encephalartos (Mogford, 1978). Heterochro-
Accepted for publication 29th August, 1977.
iit
112 Journal of South African Botany
matin has been found to be particularly frequent among the cultivated cereals such
as Triticum (Gill and Kimber, 1974), Secale and Hordeum (Vosa, 1974, 1976a),
and also among the Liliaceae such as in Scilla, Tulbaghia and Allium (Vosa, 1973
b,c, 1976 b,c).
In many such cases it has become clear that the number and position of
heterochromatin bands constitutes a character as important in the study of species
relationships as are chromosome number and morphology themselves. Equally,
however, among plant species the occurrence of heterochromatin in large amounts
remains exceptional. The discovery of large quantities of heterochromatin in the
South African Lachenalia tricolor, to be described in the present work, was
therefore considered of particular interest.
MATERIAL AND METHODS
Investigations were performed on plants from a clonal population of Lachena-
lia tricolor growing in the garden of the Department of Plant Sciences, Rhodes
University, and on plants of the variety quadricolor obtained from a commercial
source. Voucher specimens of these varieties are deposited in the Rhodes Univer-
sity Herbarium as accessions number 23793 and 23794 respectively.
In all cases rapidly growing root tips were used, pretreated with 0,1 % colchicine
for four hours and fixed in 3:1 ethanol: glacial acetic acid for 8-16 hours.
For studies of chromosome number and morphology, root tips were hydrolysed
in IN HCl at 60 °C for 6 minutes, incubated in Feulgen reagent for 90 minutes at
room temperature, and squashed in acetic orcein.
For the recognition of heterochromatin, three techniques were used: quinacrine
fluorescence (Caspersson et al., 1968); denaturation-reannealing followed by
staining with the benzimidazole derivative Hoechst 33258 (Vosa, 1976b); and
denaturation-reannealing followed by Giemsa staining (Pardue and Gall, 1970).
The procedures were as follows. For all methods except quinacrine staining,
the root tips were first hydrolysed in 0,2N HCI at 60 °C for 2,5 minutes. For all
treatments, the root tips were then squashed in 45% acetic acid under an albu-
menised coverslip, the coverslip then being floated off in absolute alcohol and air
dried. Subsequent steps were conducted on the cells adhering to the coverslip.
For quinacrine fluorescence, the preparations were stained in 0,5 % quinacrine
dihydrochloride (Gurr’s Atebrine) in absolute alcohol for 5 minutes at room
temperature. Excess quinacrine was rinsed off in absolute alcohol, and the
preparation air-dried, following which the preparation was mounted in 50 % glyce-
rine and viewed using a Zeiss fluorescence microscope with exciter filter BG12
and barrier filters 50 plus 53.
For the denaturation-reannealing treatments, the preparations after the first
ait-drying were denatured by immersion for 6 minutes in a saturated solution of
barium hydroxide at room temperature. The preparations were then rinsed tho-
roughly in running distilled water, and reannealed by immersion in 2 X saline
Centromeric heterochromatin in Lachenalia tricolor 113
sodium citrate buffer for 30 minutes at 60 °C. The preparations were then either
stained in 0,02 % alcoholic Hoechst 33258, with this and the subsequent steps as
for quinacrine staining, or else were immersed overnight in a 0,5 % aqueous solution
of Giemsa buffered to pH 6,8. The latter preparations were then air dried and
mounted permanently in Gurr’s Depex.
RESULTS
The diploid chromosome number was found to be 14, as previously deter-
mined by Spelta (1972). All the chromosomes were very small, ranging in size
from 1,1 to 2,6 uw. The karyotype was markedly asymmetric, being composed of
one, relatively large, telocentric pair (pair 1, Fig. 1), one large and two small
acrocentric pairs (nos. 3, 4 and 5), and one large and two small metacentric pairs
(nos. 2, 6 and 7).
The techniques for heterochromatin recognition indicated that heterochromatin
was present on all the chromosomes, and was exclusively centromeric in position.
The heterochromatin was uniform in its staining response, showing positive
stainability in all three techniques.
Of the various techniques, quinacrine staining was the least satisfactory, due to
the rapid fading of fluorescence. In contrast, denaturation-reannealing followed by
Hoechst staining gave a brilliant, stable fluorescence which clearly indicated the
location of heterochromatin despite the small size of the chromosomes. The results
with Giemsa were comparable but less distinct.
Chromosome number, morphology, and the position of heterochromatin were
all constant throughout both cultivars studied.
DISCUSSION
The present work emphasises further the widespread occurrence of heterochro-
matin among the Liliaceae, the significance of which is not yet clear. Stebbins
(1971) regarded heterochromatin as particularly characteristic of highly specialised
species—a viewpoint which, though fully supported by the recent demonstration
of heterochromatin in Encephalartos (Mogford, 1978), is nevertheless difficult to
apply to individual species such as the present without adequate comparative
studies within the groups concerned.
From the purely functional viewpoint, it is of interest that all the heterochro-
matin in the present species is centromeric in position. Such is the case even with
the telocentric chromosomes, which morphologically are highly dissimilar to the
remainder. Terminal and intercalary bands, so frequent in the other cases cited
above, are not in evidence. ae
Moreover, no differences occurred between chromosomes in the staining
responses of the heterochromatin. This contrasts with the situation which exists,
for example, in Allium flavum, in which different heterochromatin bands show all
114
Journal of South African Botany
A
a suisse
4 55-4
Centromeric heterochromatin in Lachenalia tricolor ILLS)
eq "ea.
wt
‘
Pa
MS
Fic. 1.
The karyotype of Lachenalia tricolor
A, Idiogram. Stippling indicates position of heterochromatin; B,
Feulgen/Acetic orcein staining, phase contrast illumination; C,
Quinacrine, fluorescence; D, Barium/SSC/Hoechst, fluorescence;
E, Barium/SSC/Giemsa, brightfield illumination.
All x 1 500.
116 Journal of South African Botany
four combinations of enhanced or reduced quinacrine fluorescence, and positive or
negative stainability with Giemsa (Vosa, 1973d).
The constant position and stainability of heterochromatin in this species
suggests that such has a uniform and specific role. Brown (1966) suggested that
heterochromatin in the vicinity of the centromeres, and also that in the vicinity of
the nucleolar organisers, might serve to separate off the highly specialised
activities of these regions from those of the remainder of the chromosome.
Nowadays, however, other possibilities may be cited. Heterochromatin, for
instance, is now known to be characterised by a highly repetitive base constitution,
which suggests that nucleolar organiser heterochromatin might be involved in
nucleolar synthesis (Mogford, 1978); while centromeric heterochromatin might
well be involved in chromosome organisation during interphase, or meiotic pairing
(Mogford, 1977). Further work is necessary to examine these possibilities.
REFERENCES
Brown, S. W., 1966. Heterochromatin. Science, N.Y. 151: 417-425.
CASPERSSON, T., FARBER, S., FOLEY, G. E., KupyNowski, J., Mopest, E. J.,
Simonsson, E., WAGH, U. and ZECcH, L., 1968. Chemical differentiation along
metaphase chromosomes. Exp! Cell Res. 49: 214-222.
DARLINGTON, C. D. and La Cour, L. F., 1940. Nucleic acid starvation of
chromosomes in Trillium. J. Genet. 40: 185-211.
Gitt, B. S. and Kimper, G., 1974. Giemsa banding and the evolution of wheat.
Proc. natn. Acad. Sci. U.S.A. 71: 4086-4090.
Marks, G. E. and SCHWEIZER, D., 1974. Giemsa banding: karyotype differences
in some species of Anemone and Hepatica nobilis. Chromosoma 44: 217-224.
Mocrorp, D. J., 1977. Chromosome associations in onion root tip nuclei. J/ S.
Afr. Bot. 43: 97-102.
Mocrorp, D. J., 1978. Nucleolar heterochromatin in Encephalartos. Jl S. Afr.
Bot. 44: 83-87.
ParpDUE, M. L., and Gatt, J. C., 1970. Chromosome localisation of mouse
satellite DNA. Science 168: 1356-1358.
Speta, F., 1972. Entwicklungsgeschichte und Karyologie von Elaiosomen an
Samen und Friichten. Naturk. Jb. Stadt Linz: 9-65.
STEBBINS, G. L., 1971. Chromosomal Evolution in Higher Plants. London:
Arnold.
Vosa, C. G. and Marcui, P., 1972. On the quinacrine and Giemsa patterns of the
chromosomes of Vicia faba. Gg. Bot. ital. 106: 151-159.
Vosa, C. G., 1973a. Acetic-orcein, a new stain for the detection of constitutive
heterochromatin in plant and animal chromosomes. Expl Cell Res. 79: 463—465.
Vosa, C. G., 1973b. Heterochromatin recognition and analysis of chromosome
variation in Scilla sibirica. Chromosoma 43: 269-278.
Vosa, C. G., 1973c. Quinacrine fluorescence analysis of chromosome variation in
the plant Tulbaghia leucantha. Chromosomes Today 4: 345-349.
Vosa, C. G., 1973d. The enhanced and reduced fluorescence bands and _ their
One e to the Giemsa patterns in Allium flavum. Nobel Symposium 23:
56-158. :
Centromeric heterochromatin in Lachenalia tricolor Lily
Vosa, C. G., 1974. The basic karyotype of rye (Secale cereale) analysed with
Giemsa and fluorescence methods. Heredity, Lond. 33: 403-408.
C. G., 1976a. Chromosome banding patterns in cultivated and wild barleys
(Hordeum spp.). Heredity, Lond. 37: 395-403.
Vosa, C. G., 1976b. Heterochromatic patterns in Allium. 1. The relationship
between the species of the cepa group and its allies. Heredity, Lond. 36: 383-392.
Vosa, C. G., 1976c. Heterochromatic patterns in Allium. 2. Heterochromatin
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VOSA,
JIS. Afr. Bot. 44 (2): 119-125 (1978)
THE PECULIAR LEAF TRICHOMES OF PETREA VOLUBILIS
(VERBENACEAE)
Cu. PUFF*
(Institute of Botany, University of Vienna, Austria)
ABSTRACT
The structure of the unicellular, heavily silicified leaf trichomes of P. volubilis which
account for the leaves’ “‘sand paper’’ structure is clarified: the hairs consist of a disc-like
base (in mature leaves frequently located in a shallow depression of the epidermis, and
therefore usually not easily recognisable) and a conical, sharp tip in its centre (considered to
be the whole hair by earlier workers). The development of these hairs is described, and
observations are included about distribution and structure of the other types of non-
glandular and glandular trichomes on the leaf blades of P. volubilis.
UITTREKSEL
DIE BESONDERSE BLAARTRIGONE VAN PETREA VOLUBILIS (VERBENACEAE)
Die struktuur van die eensellige, erg gesilisifeerde blaartrigone van P. volubilis wat die
“*skuurpapier’’ tekstuur van die blaar verklaar, word verduidelik: die hare het ’n skyfagtige
basis (in volwasse blare dikwels in vlak holtes van die epidermis geleé en dus nie maklik
uitkenbaar nie) en ’n koniese skerp punt in die middel (deur vorige werkers as die volledige
haar beskou). Die ontwikkeling van hierdie hare word beskryf en waarnemings aangaande
die verspreiding en struktuur van die ander tipes nie-klier- en kliertrigones op die blaarskyf
van P. volubilis word ingesluit.
INTRODUCTION
Petrea volubilis L.1, an attractive twining shrub indigenous to Central America
and the West Indies and frequently grown in (sub)tropical gardens of the New and
Old World, has evergreen, leathery leaves which are rough to the touch on both
surfaces (they feel like sand paper if rubbed between two fingers). This roughness
is due to the presence of peculiar hairs, about whose structure there exists a
number of quite diverse and controversial interpretations:
Vesque (1885) writes about ‘‘peculiar conical outgrowths of the outer epider-
mal wall which are completely silicified’’. Solereder (1899) observed (in addition
to glandular hairs which are not specific to Petrea, but characteristic for most
Verbenaceae) ‘‘short, conical, silicified trichomes which are embedded singly in
the thickened outer wall of the epidermis cells’’ and supports his description with
*Present address: Department of Botany and Microbiology, University of the Witwatersrand.
' The spelling Petraea often found in horticultural books and journals is inaccura
the frequently used author name Jacquin (Moldenke, 1938: 32).
te; so is
Accepted for publication 2nd September, 1977.
119
120 Journal of South African Botany
0 9 ; Fics 1 and 2.
Scanning electron micrographs of leaf blades of immature leaves. Fig. 1: adaxial surface,
Fig. 2: abaxial surface. Figs 1 and 2: x260. Explanations in the text.
A
The peculiar leaf trichomes of Petrea volubilis ( Verbenaceae) 121
the drawing of raindrop-shaped hairs sunk in the epidermis wall. Metcalfe and
Chalk (1950), without further comments, accept his diagnosis. Inamdar’s (1969)
description and drawing differ considerably from those of the earlier workers:
according to him the trichomes are silicified, conical and unicellular (although he
pictures a peculiar three-tipped hair with a number of ‘‘cavities’’ in the siliceous
deposits), and are positioned on top of the epidermis, the base of each hair
covering the area of numerous epidermal cells. Inamdar (/.c.), however, makes no
reference to the conflicting resuiis of the previous workers. Also Napp-Zinn
(1973) fails to discuss these discrepancies, so that up to now the exact structure of
these hairs is still unclear.
It, therefore, appeared worthwhile to re-examine in detail these trichomes,
which apparently are specific to P. volubilis (and probably Teijsmanniodendron
according to Metcalfe and Chalk, 1950) and unique in the Verbenaceae—a family
characterised by a wide range of different hair forms (cf. Briquet, 1897;
Solereder, 1899; Metcalfe and Chalk, 1950).
MATERIAL AND METHODS
Leaf material (usually 5 immature and 5 mature leaves) of P. volubilis growing
in the gardens in and around Pietermaritzburg was collected and then immediately
investigated under the light microscope (transverse sections by hand and epidermis
peels) to ensure that the trichomes in question are a stable, always present feature
of the species.
SEM investigations: Fresh leaf segments were coated with gold-palladium and
viewed with a Hitachi SSM 2 at an accelerating voltage of 10 kV. It was not
necessary to use the critical point drying method: no differences whatsoever were
observed when pictures of fresh and critical point dried leaves were compared.
TEM investigations: The material was fixed for 2 h in 3% glutaraldehyde
followed by 1 h in 1,0% 0s0,, both in 0,1 phosphate buffer (pH 7,0). After
dehydration with graded ethanol the material was embedded in Epon 812.
Ultrathin sections were post-stained with uranyl acetate and lead citrate. The
electron micrographs were obtained with a Hitachi HU 11.
OBSERVATIONS
The following types of trichomes are found on the leaf blades of P. volubilis:
(1) glandular hairs with a (A) short (one-celled) or (B) long stalk and a capitate
gland consisting of two to several (6 to 9) radially arranged secretory cells (Figs 1
and 2), and (2) non-glandular trichomes in the form of (A) + long, unicellular
hairs surrounded by a conspicuous “‘ring’’ of cells at the base (Fig. 2) and (B)
‘short, conical, unicellular hairs’’ (Solereder, /.c.) specific to the species. The
occurrence of the various hair types is different on the ad- and abaxial surface of
the blades, and also depends on the age of the leaf (Moldenke, 1938, already
122 Journal of South African Botany
mentions that the blades of mature leaves are very rough to the touch on both
surfaces, whereas those of very immature ones are less conspicuously so).
The long-stalked glandular hairs are only found on the adaxial surface of very
immature leaves, where they may occur in large numbers (and much more closely
spaced than in the leaf portion shown in Fig. 1). They seem to completely vanish
in old material (due to breaking or falling off?).
The short-stalked glands are very common on the abaxial surface of young and
old leaves, but are often absent from the adaxial surface (if present, they never
occur in large numbers and are sunk in crypts; their appearance is similar to that of
sunken glandular hairs of Avicennia marina: Fahn and Shimony, 1977: Plate 1C).
In mature leaves the number of secretory cells per gland is (6)8(9), in young and
very young leaves it appears to be less (2-6; even recognisable in Fig. 2: the
depressions on the glands correspond with the walls between two secretory cells).
Obviously the fewer celled glandular trichomes represent younger stages of
development (cf. Fahn and Shimony, 1977: Fig. 2J—N).
The very sparse, long, unicellular hairs (Figs 1 and 2) are found on the abaxial
surface (particularily in young material) and (very seldom) on the upper epidermis,
where they seem to completely disappear in mature leaves.
| 7 a
Fics 3 and 4.
Scanning electron micrographs of adaxial surface of mature leaves. Fig. 3: x260 Fig. 4:
400. Explanations in the text.
The peculiar leaf trichomes of Petrea volubilis (Verbenaceae) 123
Fics 5 and 6.
Transmission electron micrographs of unicellular hairs. Fig. 5: off-median section of young,
not yet silicified hairs (immature leaf), Fig. 6: close-to-median section of fully developed,
heavily silicified hair (mature leat); silicifications partially broken out. Nu: nucleus, Si:
siliceous deposits, tW: thickened, layered cell wall, Va: vacuole. Figs 5 and 6: 4200.
124 Journal of South African Botany
The ‘‘short, conical hairs’’ sensu Solereder (/.c.) are densely distributed on
both surfaces, although usually in much greater abundance on the adaxial surface
(compare Figs 1 and 2). Although already present in immature leaves (Figs | and
2) they become particularily conspicuous on the adaxial surface of mature leaves
(Figs 3 and 4), where they, in fact, look like Vesque’s (/.c.) ““peculiar outgrowths
of the epidermis’’. Contrary to Solereder’s (J.c. ) interpretation, however, they are
not raindrop-shaped and embedded in the outer epidermal wall: longitudinal
sections prove that they actually sit singly on top of cells and, strictly speaking,
are not conical, but consist of a flat, more or less disc-like base with a conical tip
in its center, both of which are, in the mature stage, heavily silicified (Fig. 6).
Solereder (/.c.) considered these tips, which are responsible for the ““sand paper
texture’’ of the plant’s leaves, to be the whole hairs.
iG, Vo
Schematic representation of three stages of development of the peculiar, unicellular
trichomes. A: enlarged epidermis cell (hair initial), B: young hair with starting silicification,
C: mature, heavily silicified hair. Nu: nucleus, Si: siliceous deposits, Va: vacuole. Further
explanations in the text.
The development of these trichomes is as follows (Fig. 7): at an early stage of
development of the leaf an epidermis cell increases in size and extends above the
surrounding epidermal cells (Fig. 7A). This hair initially shows no significant
differences from the neighbouring cells (cytoplasm dense, occupying almost the
whole cell lumen; few, small vacuoles; large nucleus). After a periclinal and
somewhat unequal division, hair cell and basal cell originate. The basal cell is
usually larger than the hair cell and is generally not distinguishable from the
neighbouring epidermis cells (Fig. 7B). The protoplast of the basal cell soon forms
a layer against the cell wall so that a large central vacuole originates, while the
cytoplasm still occupies most of the cell lumen of the hair cell (Fig. 5). It is
presumed that, at this stage at the latest, the silification of the conical tips of the
hairs begins, although there is no concrete evidence for this (median sections of
this stage of development always ripped out in the region of the tips indicating that
it was already filled with difficult-to-cut siliceous deposits). As development
continues the walls of the basal cells and the neighbouring epidermis cells thicken
rapidly (the outer epidermal walls more than the inner ones) and become
conspicuously layered (Fig. 6), and the hairs (whose walls do not thicken to that
extent) become almost completely filled with siliceous deposits (Figs 6 and 7C).
The peculiar leaf trichomes of Petrea volubilis (Verbenaceae) 125
At this stage (i.e. in mature leaves), the outer epidermal walls have become quite
uneven, and the disc-like bases of the hairs often are somewhat obscured, since
they are no longer—as in immature leaves—clearly raised above the level of the
outer epidermal walls of the surrounding cells (compare Figs 1, 2 and 3, 4 and 6).
ACKNOWLEDGMENTS
The present investigation was carried out while at the Department of Botany,
University of Natal, and I would like to thank Mrs M. G. Gilliland, Officer-
in-charge of the Electron Microscope Unit, and Mr V. J. Bandu for their
invaluable assistance in electron microscopy and the preparation of the photogra-
phical prints.
REFERENCES
BRIQUET, J., 1897. Verbenaceae. In: A. Engler und K. Prantl: Die natiirlichen Pflanzen-
familien. Band IV. 3a: 132-182.
FaHN, A. and SHIMONy, C., 1977. Development of the glandular and non-
glandular leaf hairs of Avicennia marina (Forsskal) Vierh. J. Linn. Soc., Bot 74:
37-46.
INAMDAR, J. A., 1969. Epidermal structure and ontogeny of stomata in some
Verbenaceae. Ann. Bot. 33: 55-66.
Metcatre, C. R. and CHALK, L., 1950. Anatomy of the Dicotyledons. Vol. 2.
Oxford: Clarendon Press.
Mo.LpENKE, H. N., 1938. A monograph of the genus Petrea. Feddes Rep. 43:
1-48, 161-221.
Napp-ZINN, K., 1973. Anatomie des Blattes. Il. Blattanatomie der Angiospermen.
Berlin und Stuttgart: Borntraeger.
SOLEREDER, H., 1899. Systematische Anatomie der Dicotyledonen. Stuttgart:
nke.
Vesque, J., 1885. Caractéres des principales familles gamopétales, tires de
l'anatomie de la feuille. Ann. Sci. nat. Je Sér., Bot. 1: 183-360.
re
ny a Re
vit
t:
MARY GUNN LIBRARY
NATIONAL BOTANICAL INSTITUTE
PRIVATE BAG X 101
PRETORIA 0001
REPUBLIC OF SOUTH AFRICA
JIS. Afr. Bot. 44 (2): 127-141 (1978)
LEAF ANATOMY OF JUBAEOPSIS CAFFRA BECC.
B. L. ROBERTSON
(Department of Botany, University of Port Elizabeth)
ABSTRACT
The anatomy of the lamina and leaf sheath of Jubaeopsis caffra, an endemic palm from
the east coast of South Africa, was studied. Observations were recorded and comparisons
were made with other cocosoid species. Generally the leaf anatomy of J. caffra was found
to be similar tc that of most other cocosoid genera, but a few distinguishing differences
were noted. The most significant of these is in respect of the anticlinal walls of the adaxial
epidermis cells. These cell walls are sinuous and distinguish Jubaeopsis from all cocosoid
genera except Orbignya. Further, the margins of the leaflets are slightly ribbed. This is in
contrast to the other cocosoid species where no marginal rib is present.
UITTREKSEL
BLAARANATOMIE VAN JUBAEOPSIS CAFFRA BECC.
Die anatomie van die lamina en blaarskede van Jubaeopsis caffra, ’n endemiese palm
vanaf die Suid-Afrikaanse ooskus, is ondersoek. Waarnemings word beskryf en ’n verge-
lyking met ander cocosoide spesies word gemaak. Oor die algemeen stem die blaaranatomie
van J. caffra ooreen met die meeste ander cocosoide genera, maar ’n paar uitstaande
kenmerke is waargeneem. Die belangrikste hiervan is ten opsigte van die antiklinale wande
van die adaksiale epidermisselle. Hierdie selwande is gekartel en onderskei Jubaeopsis van
alle ander cocosoide genera behalwe Orbignya. Verder, is die rande van die blare effens
gerib. Dié is in teenstelling met die ander cocosoide spesies waar geen marginale rib
aanwesig is nie.
INTRODUCTION
Tomlinson’s (1961) classic work on the anatomy of the palms deals so
completely with the literature on this subject that it would be superfluous to give a
complete review thereof here. Although Tomlinson does describe the anatomy of
the cocosoid palms in general and that of a number of genera in particular, no
information concerning the anatomy of Jubaeopsis caffra is available. In view of
this it was felt that a study of the anatomy of this species would be justified.
Although the leaf, stem and root were examined, it is the anatomy of the leaf,
and in particular that of the lamina, which exhibits by far the most diagnostic
features (Tomlinson, 1961). Consequently only data relating to the anatomy of the
lamina and leaf sheath are presented in this paper.
MATERIAL AND METHODS .
All the plant material used in this study was collected from trees in the Jubaeopsis
groves on the estuary of the Mtentu river in Pondoland on the east coast of South
Accepted for publication Sth October, 1977.
127
128 Journal of South African Botany
Africa. Fixation of the material was in formaldehyde/ethyl alcohol/acetic acid
(Sass, 1958).
Dehydration was accomplished with a graded tertiary butyl alcohol (TBA)-
ethyl alcohol series while infiltration with wax was achieved by using a graded
TBA-paraffin wax series. The material was embedded in paraffin wax (Carowax;
melting point 54-58 °C).
Sections were cut on a rotary microtome at a thickness of 10 wm. These were
stained with safranin and counterstained with fast green (Brooks, Bradley &
Anderson, 1950; Holtzhausen, 1972).
Material used in the SEM study was air-dried and coated with palladium-gold
(Robbertse, 1974) prior to viewing in a Jeol JSM-U3 microscope.
Although some sections of Jubaea material were made, the comparisons of
Jubaeopsis with Jubaea and other species are discussed mainly on the basis of
Tomlinson’s (1961) descriptions of these species.
RESULTS
Lamina
The mesophyll is clearly demarcated into adaxial palisade tissue and abaxial
spongy parenchyma and consequently the leaf of J. caffra is dorsiventral (Fig. 1).
Fic. 1.
Longitudinal section of the leaflet lamina of J. caffra. x 150.
(Ad—adaxial epidermis; h—hypodermis).
Thin flakes or layers of wax occur on the surface of the leaflets (Fig. 2) but
only a small amount of wax is present.
Epidermal hairs also occur on both surfaces. These hairs consist of a larger,
thin-walled sunken basal cell and four or five thin-walled distal cells arranged
radially around the distal tip of the basal cell (Figs 3 & 4). These hairs are
restricted to the intercostal regions. A second type of epidermal appendage occurs
in the form of scale-like outgrowths along the midrib.
Leaf anatomy of Jubaeopsis caffra Becc. 129
Fic. 2.
Stereoscan micrograph of stoma. Xx 1000.
(gc—guard cell; lcc—lateral contact subsidiary cell; lsc—tlateral subsidiary cell; tsc—
terminal subsidiary cell).
The adaxial surface is covered by a thick cuticle while that on the abaxial
epidermis is slightly thinner (Fig. 1). The cuticle is smooth and has no characteris-
tic features or patterns, except on the guard cells. (These cells are discussed later).
Along the leaflet margins and on the adaxial side of the midrib the cuticle is
particularly thick (Figs 6, 7 & 10).
In J. caffra the outer tangential walls of the epidermis cells are largely
cutinised and are thick (Figs 1, 3, 6-8, 10, 11) especially in the case of the adaxial
epidermis and in the vicinity of the midrib and margin. The cell walls are however
not lignified. While the tangential or anti-clinal cell walls of the adaxial epidermis
appear somewhat sinuous in a surface view (Fig. 5) those of the abaxial epidermis
are only slightly sinuous to non-sinuous (Figs 12 & 13).
Adaxially no costal and intercostal bands are distinguishable, but abaxially the
epidermis cells of these two regions differ mainly in respect of size with the costal
cells slightly larger and broader than the intercostal epidermis cells (Fig. 12). The
costal bands are narrower than the intercostal ones (Fig. 12).
The cells of the epidermis are longitudinally extended but are rather shallow
(Figs 1, 3, 11 & 12). Around the leaf margin and the midrib however, the
130 Journal of South African Botany
Longitudinal section through a portion of the lamina showing an adaxial epidermal hair
(eph). X 150.
Fic. 4.
Surface view of the epidermal hair (eph). 400.
1G, De
Surface view of the adaxial epidermis of the leaf. x 180.
Fics 6 & 7.
Transverse section through leaflet midrib. x60; x150 resp. (ad—adaxial surface; mc—
motor or expansion cells; mx—metaxylem; p—peripheral bundle; ph—phloem, sch—
sclerenchyma; sp—sclerotic parenchyma).
Fic. 8.
Transverse section through lamina. x 150. (ad—adaxial surface; fs—fibrous strands; mx—
metaxylem; ph—phloem; ps—parenchymatous sheath; px—protoxylem; rs—raphide sac;
s—stoma; ss—sclerenchymatous sheath; svb—small vascular bundle).
Leaf anatomy of Jubaeopsis caffra Becc. Bil
epidermis cells are uniformly isodiametric (Figs 7 & 10). The abaxial epidermal
cells over the hypodermal expansion or motor cells on either side of the midrib are
distinctly papillose (Fig. 6).
The stomata are restricted to the abaxial surface and specifically to the
intercostal bands where they are very numerous (Fig. 12). The stomatal complex
found in this species appears to be similar to that described by Tomlinson (1974)
for Cocos and other Palmae, viz. a complex which originates from non-
intersecting oblique divisions in the neighbouring cells. Each stoma is surrounded
by six subsidiary cells (Figs 2 & 13). Two of these are terminal subsidiary cells and
are short, thin-walled kidney-shaped cells (Figs 2 & 13) which are not sunken
(Fig. 2). The two lateral contact cells are very narrow cells with the non-contact
walls, i.e. walls not in contact with the guard cells being thickened (Fig. 13).
These cells are distinctly sunken and envelope the guard cells (Figs 2, 8 & 14).
The remaining two large lateral subsidiary cells are not sunken (Figs 2 & 14). The
guard cells, which are sunken (Figs 2, 8 & 14) are small and thin-walled and are
provided with a prominent outer and slightly less prominent cutinised ledge (Fig.
14).
A hypodermis of large cells occurs beneath both the adaxial and abaxial
surfaces. In each case only a single layer of hypodermal cells occurs (Fig. 1 & 8)
except in the vicinity of the margin and the midrib (Figs 6, 7, 10 & 11). Cells of
the hypodermal layers are thin walled. The shape and size of the adaxial and the
abaxial hypodermal cells tend to be irregular (Figs 8 & 12). In the abaxial
hypodermis the intercostal hypodermis cells are somewhat larger than the costal
hypodermis cells (Figs 8 & 12).
Each substomatal chamber is surrounded by four L-shaped abaxial hypodermis
cells (Fig. 9).
( 10
Fic. 9.
Paradermal section through abaxial hypodermis of the leaf showing four L-shaped cells (1-4)
around the sub-stomatal chamber. <360.
if” @
Fic. 10. :
Transverse section through the leaf margin. x90. (ad—adaxial surface; css—connecting
strip scar.)
132 Journal of South African Botany
The chlorenchymatous mesophyll is clearly demarcated into an adaxial pali-
sade tissue and abaxial spongy parenchyma (Fig. 1).
Generally only one or two layers of palisade cells are present. In a surface
view the palisade cells which appear more or less circular, are densely packed into
neat longitudinal files between the strands of fibres which run parallel to the long
axis of the leaflet (Fig. 15), while the cells of the spongy parenchyma are extended
transversely (Fig. 16) at right angles to the axis of the leaflet.
Three types of idioblasts occur in the mesophyll, viz. (i) a large number of
tanniniferous cells, (1i) raphide-sacs and (iii) silica cells or stegmata. The tannini-
Longitudinal section through leaflet midrib. x 150. (ad—adaxial surface; h—hypodermis;
sf—septate fibres).
Fic. 12.
Oblique surface section through the abaxial epidermis of the lamina. x60. (cb—costal band;
ep—abaxial epidermis; h—hypodermis; icb—intercostal band; sp—spongy parenchyma).
Fic. 13.
Surface view of the abaxial surface of the lamina showing the organisation of the stomatal
components. 360. (gc—guard cells; 1cc—lateral contact cell; Isc—later subsidiary cell;
tsc—terminal subsidiary cell).
Fic. 14.
Transverse section through a stoma. X360. (cl—cuticular ledge; lcc—lateral contact cell;
gc—guard cells; lsc—tlateral subsidiary cell; P—pore; stc—substomatal chamber).
Leaf anatomy of Jubaeopsis caffra Becc. 133
ie GID
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Fic. 15.
Paradermal section through the lamina of a leaflet, showing the closely packed
chlorenchyma and fibrous strands. x 160. (fs—fibrous strands; pc—palisade chlorenchyma).
Fic. 16.
Paradermal section through the abaxial portion of the lamina, showing the transversally
extended spongy parenchyma cells. 160. (ics—intercellular spaces; sp—spongy paren-
chyma).
sheath. Sacs containing raphide clusters occur as large cells elongated parallel to
the long axis of the leaflets (Figs 8 & 17). Each cell contains a bundle or sheaf of
needle-like crystals which are embedded in densely staining mucilage. With regard
to the stegmata, the silica bodies in cells are spherical. Although these cells are
numerous, they are found only in association with vascular bundles and fibrous
bundles. The stegmata occur in single longitudinal files around these bundles (Fig.
18). The walls of the stegmata are slightly lignified and very often these cells
occur in depressions in the walls of the fibres.
Numerous non-vascular fibres are present in the lamina. These occur singly
and in groups or strands and are situated mainly beneath the adaxial hypodermis
(Fig. 8). Only a small number of fibrous strands occur abaxially. In a transverse
section these fibrous strands are small and consist mainly of between one and six
fibres. At the margin however, the fibrous bundles are much thicker (Fig. 10).
These fibres are thick-walled, septate (Fig. 15) and often unlignified.
The venation of the lamina is parallel and includes longitudinal veins of
various sizes connected by transverse commisures (Fig. 19). Basically two sizes of
veins occur (excluding the midrib). The largest of these, which often contains a
single large metaxylem vessel, is more or less centrally situated in the mesophyll
and is connected to the epidermis by a fibrous sheath. The small vascular bundles
on the other hand are restricted to the abaxial mesophyll and connected to the
abaxial surface only (Fig. 8).
The large vascular bundles are sheathed by an outer parenchymatous layer and
an inner sclerotic layer. The outer sheath is uniseriate and consists of large,
134 Journal of South African Botany
6 Oak asl Be
lG, IW,
Paradermal section through the lamina showing a raphide sac (rs) in the mesophyll. 230.
Fic. 18.
Longitudinal section through lamina showing stegmata and silica bodies associated with
fibres. < 570. (ffibres; st—stegmata with spherical silica bodies).
vertically extended (in a transverse section) parenchyma cells containing only a
small number of chloroplasts. This sheath usually occurs only laterally to the vein
and is often absent from the adaxial and abaxial regions (Fig. 8). The inner sheath
is multiseriate and is comprised of fibres and sclerotic parenchyma, the fibres
being limited mainly to the adaxial zone (Fig. 8). These fibres are lignified and are
easily distinguishable from the adaxial fibre buttresses which connect the veins
with the adaxial hypodermis. The outer sheath of the small veins is not well
differentiated. The inner sheath is also somewhat reduced and consists adaxially of
a single layer of sclerotic parenchyma and abaxially of two or three layers of
fibres.
The xylem consists mainly of protoxylem vessels and xylem parenchyma. It is
normally very difficult to distinguish between these tissues and the sclerenchyma-
tous sheath, especially in the small veins, when viewed in transverse section. The
Leaf anatomy of Jubaeopsis caffra Becc. 135
larger veins however normally contain a single large metaxylem vessel which is
easily distinguishable (Fig. 8).
The phloem occurs abaxially in the vascular bundles. It is separated from the
xylem by at least one cell layer of sclerenchyma (Fig. 8). Normally the phloem,
which includes sieve tubes, companion cells and phloem parenchyma is divided
into more than one group in the midrib only, but not in the other veins (Figs 6-8
& 10).
The midrib is adaxially very prominent (Fig. 6). As mentioned earlier, the
epidermis cells are fairly uniform, but the abaxial ones are somewhat smaller than
the adaxial epidermis cells, and the adaxial cuticle is thicker than that of the
abaxial surface (Fig. 7).
Adaxially the hypodermis of the midrib is multiseriate and comprises three
layers of large parenchyma cells (Figs 6, 7 & 11). Abaxially, the hypodermis cells
are small in the centre of the rib, but are laterally confluent with the expansion
cells and thus increase in size on either side of the midrib (Figs 6 & 7).
In the midrib there is a thick multiseriate fibrous cylinder surrounding the
central ground tissue. This cylinder is composed of two crescent-shaped sections,
one adaxially and the other abaxially orientated. Where these two sections join,
i.e. more or less across the middle of the midrib (in transverse section) the cell
walls are somewhat thinner (Figs 6 & 7).
The central ground tissue, which consists mainly of sclerenchymatous tissue or
sclerotic parenchyma, envelops the vascular bundles. A large vascular bundle is
situated abaxially and consists of xylem (with a single large metaxylem vessel)
—— Seer Se Dee
Be i wen 5
: PAR GAY Sela BEE, if
—
pes
AS aS OR
Fic. 19.
Paradermal section of leaflet in which transverse commisures are evident. x25. (cb—costal
bands; icb—intercostal bands; tc—transverse commisures).
Fics 20 & 21. Se
Transverse section through the dorsal portion of the spear-leaf sheath. eons ei
(mx—metaxylem; pg—starch-rich parenchymatous ground tissue, ae nen Pp
protoxylem; ufs—undifferentiated fibrous sheath or cap, vt—vascular tissue).
136 Journal of South African Botany
and three or four small groups of phloem separated from each other and from the
xylem by partitions of sclerenchyma. A second, smaller vascular bundle is
adaxially situated, but is of a more compact nature (Figs 6 & 7). Peripheral
vascular bundles do occur in the midrib (Fig. 7) but are rare.
Two abaxial bands of expansion cells occur in the leaflet. These are present as
bands on either side of the midrib and run parallel to the midrib (Fig. 6). They
consist of one or two hypodermal layers of very large, colourless parenchyma cells
which occupy more than half the depth of the lamina. Although the two bands are
confluent with the hypodermis of the midrib the cells of the midrib hypodermis are
not differentiated into expansion cells. No fibrous strands occur within these
bands.
Although the lamina of cocosoid palms does not normally have a well-
differentiated margin, that of J. caffra is rather prominent (Fig. 10). The cuticle,
epidermis and hypodermis in this region are very similar to those of the midrib.
Further, the veins in the margin possess very thick abaxial fibrous caps. The
epidermis of the lamina is interrupted at a point near the margin by the “‘scar”’
left by the connective strip which connects the leaflets of the lamina with each
other in the developing leaf (Fig. 10).
Leaf sheath
The sheath of the spear leaf, i.e. the oldest, unopened leaf, is still meristematic
and consequently it is largely parenchymatous and no sclerotic tissue has differen-
tiated at this stage (Fig. 20). However, associated with each vascular bundle is a
group of cells which are much smaller in diameter than the cells of the
surrounding parenchymatous ground tissue (Figs 20 & 21).
These cells constitute undifferentiated fibres which, after differentiation, give
the sheath its tremendous strength and durability. The peripheral bundles in the
dorsal section of the sheath are associated with exceptionally large, fibrous caps
(Fig. 20). In the spear leaf the vascular tissue in the leaf sheath consists of
protoxylem, one group of phloem per bundle and vascular parenchyma. Differen-
tiation of metaxylem is initiated at about this time (Fig. 21).
The cells of the ground tissue are very rich in starch (Figs 21 & 23).
In the sheath of the youngest expanded leaf, the cells are also still mainly
unlignified and no sclerenchyma is evident (Figs 22-25). The adaxial cells of the
ventral section however, become tangentially elongated, probably as the result of
the expansion in diameter of the stem (Fig. 23). Apart from the group of
undifferentiated fibres associated with the vascular bundles, numerous strands of
undifferentiated non-vascular fibres become evident. Although these are scattered
throughout the ground tissue (Figs 22 & 25) they are concentrated in the abaxial
peripheral region (Fig. 25). At this stage each vascular bundle consists of a large
undifferentiated fibrous sheath, a number of protoxylem vessels, a single large
metaxylem vessel, two groups of phloem and vascular parenchyma (Fig. 24).
Leaf anatomy of Jubaeopsis caffra Becc. 13)7/
oe
res a
ih?
satan at wits
Fics 22-25.
Transverse section through the ventral portions of the sheath of the youngest expanded leaf.
Fic. 22.
SOS,
EIGs 23"
Portion B of Fig. 22. 160.
Fic. 24.
Portion C of Fig. 22. x 160.
EIG@25:
Portion D of Fig. 22, showing the peripheral fibrous bundles (still unsclerified) in the
abaxial portion of the leaf sheath. x 160.
(ab—abaxial surface; ad—adaxial surface; f—‘‘filling’’ fibrous strands; mx—metaxylem,
Pg—parenchymatous ground tissue; ph—phloem; px—protoxylem; ufs—unsclerified fibrous
strand; ufsh—unsclerified fibrous sheath; wp—‘‘warp’’ vascular bundles; wt—‘‘weft’’
vascular bundles).
As the leaf matures, sclerification of the fibres takes place and each vascular
bundle is associated with a large sclerotic cap (Fig. 26). The number of metaxylem
elements increases, but still only two groups of phloem are present. Further, a
large amount of sclerotic parenchyma occurs in the vascular bundles (Fig. 27). In
a longitudinal section through the ventral tissues of the sheath, the vascular
bundles are not perfectly vertical but are arranged at a slight angle to the left and
right. The adaxial bundles which constitute the ‘‘weft’’, are angled upwards from
right to left on the one side of the ventral line and from left to right on the
opposite side of this line. The abaxial bundles, i.e. the ‘‘warp’’ on the other hand,
are orientated in precisely the opposite manner. The type of leaf sheath found in
138 Journal of South African Botany
0 cpg,
. ae 6 GES
ee
4 “i
per Spa
See
ao
Fic. 26.
Transverse section through the ventral tissues of the sheath of a mature expanded leaf,
showing the extent to which the fibrous sheath or phloem cap becomes sclerified. 25.
(scl—sclerenchyma).
Ine, ZI,
Enlargement of one of the vascular bundles in Fig. 26. x160. (mx—metaxylem; ph—
phloem; px—protoxylem; scl—sclerenchyma; sp—sclerotic parenchyma).
Fic. 28.
Transverse section through the abaxial ventral tissues of a mature leaf sheath showing the
““warp’’ vascular bundles. x25.
Fic. 29.
Transverse section through the adaxial vascular bundles of the same leaf as in Fig. 28.
These bundles constitute the ‘‘weft’’. x25.
Leaf anatomy of Jubaeopsis caffra Becc. 139
Jubaeopsis caffra is intermediate between the Phoenix and Cocos types (Robert-
son, 1977). As can be seen from Figs 28 and 29, there is very little difference in
thickness and the extent of sclerification between the warp and weft bundles. The
numerous, non-vascular fibrous bundles constitute the ‘‘filling’’. (The terminology
used here is that proposed by Tomlinson, 1964).
DISCUSSION AND CONCLUSIONS
The dorsiventrality of the leaf of J. caffra is in accordance with the majority of
other cocosoid genera.
The epidermal hairs which occur on the leaf of J. caffra are similar to those of
Orbignya, Elaeis and Syagrus in that they consist of a single inflated, distinctly
sunken basal cell and a number of thin-walled distal cells, but differ from those of
Jubaea in which the basal cell is not sunken and from those in Cocos where three
or four basal cells occur.
As far as the adaxial epidermis of the leaf is concerned, J. caffra differs from
all the cocosoid palms except Orbignya, in that the anticlinal cell walls are
sinuous. Further, J. caffra differs markedly from Jubaea in that the adaxial
epidermis cells of the former species are longitudinally elongated, while in the
latter case, the cells are transversely elongated (Tomlinson, 1961).
As in other cocosoid genera, the stomata of J. caffra are restricted to the
intercostal bands of the abaxial surface. The stomatal complex of J. caffra is
similar to that described by Tomlinson (1974) for Cocos nucifera. According to
Tomlinson (1961) the guard cells are not sunken, except in Scheelea and Syagrus
yatay. In this respect J. caffra is similar to the latter two cases in that the guard
cells are distinctly sunken (Figs 2 & 13).
Both the adaxial and abaxial hypodermis layers of J. caffra are only one cell
layer thick (Fig. 1). In this respect it is similar to Elaeis and Orbignya, but
dissimilar to Cocos which has an adaxial hypodermis of one or two cell layers.
Jubaea on the other hand has a well-developed adaxial hypodermis, consisting of
three cell layers while a single cell layer occurs beneath the abaxial epidermis.
The chlorenchyma of most cocosoid palms is usually demarcated into distinct
adaxial palisade layers and abaxial spongy parenchyma. J. caffra \eaves exhibit
the same structure and differ thus from Jubaea in which no spongy parenchyma
occurs. The entire mass of chlorenchyma in the leaf of this latter genus consists of
palisade tissue (Tomlinson, 1961).
Orientation of the fibrous strands in the leaf of J. caffra is similar to the
majority of genera in this sub-family in that they occur mostly as adaxial,
sub-hypordermal strands with only a few abaxial strands. Again, J. caffra differs
quite markedly from Jubaea which has very few non-vascular fibrous strands.
Similarly, J. caffra differs from Jubaea but is basically similar to the majority
of cocosoid genera in respect of the orientation of the veins in the lamina. Generally
the large vascular bundles in most genera are centrally placed in the mesophyll
140 Journal of South African Botany
and have contact with both surfaces, while the smaller vascular bundles are
abaxially situated and are connected to the abaxial surface only (Fig. 8). In Jubaea
(Tomlinson, 1961) on the other hand, all the vascular bundles, irrespective of size,
are in contact with both surfaces via fibrous buttresses.
In J. caffra three to four groups of phloem occur in the larger vascular
bundles. These are separated from each other by strips of sclerotic parenchyma
which radiate from the metaxylem vessel. In this respect it is similar to Cocos,
Coroza, Attalea, Elaeis and Maximiliana (Tomlinson, 1961) but differs from
Jubaea in which the number of phloem groups often exceeds four and which are
separated by both vertical and horizontal sclerotic partitions.
The most significant difference between the midrib of J. caffra and that of
Jubaea is to be found in the chlorenchyma. In Jubaea the central sclerotic cylinder
of the midrib is surrounded by well-developed peripheral chlorenchyma which is
continuous with that of the lamina. However, in J. caffra very little assimilatory
tisssue is present in this region (Fig. 7).
It appears that all cocosoid genera (except Maximiliana) have two abaxial
bands of expansion cells—one on either side of the midrib of the leaflet. The
composition of these bands though, is not the same in all genera. In J. caffra, for
example, the bands consist of between one and two cell layers only (Fig. 7) while
in Jubaea the equivalent cells are smaller, but between three and four cell layers
are present.
Although Tomlinson (1961) states that the margins of the leaflets in cocosoid
genera are not ribbed, J. caffra has a rather prominently developed margin,
consisting of two to three hypodermal cell layers, two vascular bundles and a
fibrous bundle. In this respect J. caffra is quite similar to the caryotoid palms
which have distinct marginal ribs (Tomlinson, 1961).
The only detailed description of a cocosoid leaf sheath is of Cocos nucifera
(Tomlinson, 1964). It would seem from the results of this study that the anatomy
of J. caffra’s sheath is very similar to that of C. nucifera, except that there is little
difference between the adaxial and abaxial bundles, i.e. weft and warp respec-
tively, in J. caffra.
The general conclusions that can be drawn from this study are that although
the anatomy of J. caffra is basically similar to the majority of other cocosoid
palms, it does have certain distinguishing features. The most important of these
being the sinuous anticlinal walls of the adaxial epidermis cells and the presence
of a marginal rib.
Anatomically, the leaf of Jubaeopsis differs very markedly from that of Jubaea
and the relationship between these two monotypic genera is perhaps not as close
as suggested by Beccari (1913).
Leaf anatomy of Jubaeopsis caffra Becc. 141
ACKNOWLEDGEMENTS
This study was supported by financial grants from the University of Port
Elizabeth and the CSIR.
REFERENCES
BeEccaRI, O., 1913. Una nuova “‘Cocoinea’’ africana: Jubaeopsis caffra. Webbia 4:
169-176.
Brooks, R. M., BRADLEY, Muriel V. and ANDERSON, Thelda I., 1950. Plant
microtechnique manual. Davis: University of California.
HOLTZHAUSEN, L. C., 1972. ’n Morfo-genetiese en fenologiese studie van die
blom en vrug van Citrus sinensis (L.) Osbeck., cultivar Valancia. D.Sc. (Agric.)
thesis. Pretoria: University of Pretoria.
RopBerTsE, P. J., 1974. A scanning electron microscopic investigation of the
pollen of South African Acacia species. JI S. Afr. Bot. 40: 91-99.
ROBERTSON, B. L., 1977. Aspects of the morphology of Jubaeopsis caffra Becc.
Accepted for publication in October issue of Principes.
Sass, J. E., 1958. Botanical microtechnique. lowa: lowa State University Press.
ToMLINSON, P. B., 1961. Anatomy of the monocotyledons, WU. Palmae. London:
Oxford University Press. 5:
ToMLINSON, P. B. 1964. The vascular skeleton of the coconut leaf base.
Phytomorphology 14: 218-230.
ToMLINsON, P. B., 1974. Development of the stomatal complex as a taxonomic
character in the monocotyledons. Taxon 23(1): 109-128.
JIS. Afr. Bot. 44 (2): 143-151 (1978)
A PRELIMINARY STUDY OF THE NITROGEN NUTRITIONAL STATUS
OF MEMBERS OF THE SOUTH AFRICAN PROTEACEAE
O. A. M. Lewis and W. D. Stock
(Department of Botany, University of Cape Town)
ABSTRACT
An investigation of the free amino compounds of Leucadendron xanthoconus and
Protea lepidocarpodendron leaves indicate very low concentrations of these compounds
present. An analysis of the xylem sap of L. xanthoconus, P. lepidocarpodendron, P.
laurifolia and Brabeium stellatifolium also reveals very low levels of nitrogen transporting
compounds present, the spectrum of these compounds being dominated by nitrate and
ammonium ions, and glutamine. Feeding experiments with KNO3 and NH,CI indicate poor
utilisation of additional nitrogen supplies by L. xanthoconus shoots, particularly in the case
of nitrate feeding. '"N feeding experiments do, however, show limited nitrate reduction
capability in the leaves of L. xanthoconus.
UITTREKSEL
*N VOORLOPIGE STUDIE VAN DIE STIKSTOF-VOEDINGSSTATUS VAN LEDE
VAN DIE SUID-AFRIKAANSE PROTEACEAE
*n Ondersoek van die vrye aminoverbindinge van Leucadendron xanthoconus en Protea
lepidocarpodendron blare vertoon die aanwesigheid van baie lae konsentrasies van hierdie
verbindinge. Oplossings van die xileemsap van L. xanthoconus, P. lepidocarpodendron, P.
laurifolia en Brabeium stellatifolium besit ook ’n baie lae konsentrasie van stikstofvervoer-
verbindinge, met nitraat- en ammoniakione, en glutamien as die hoofkomponente. Voedings-
eksperimente met KNO, en NH,C1 vertoon swak benutting van sulke bykomende stikstof-
bronne deur uitspruitsels van L. xanthoconus, veral in die geval van nitraatvoeding. *N
voedingseksperimente het egter bewys dat die blare van L. xanthoconus oor ’n beperkte
vermoe beskik om nitraat te reduseer.
Although the fynbos of the South Western Cape has been known to civilised
man longer than any other vegetation type in Southern Africa, it remains one of
the most poorly studied of these types, especially in relation to its ecophysiologi-
cal relationships. This apparent neglect of the country’s most spectacular flora is
probably occasioned by three main factors: the complexity of the flora itself; the
fact that the topography and soils outside the regions exploited early in the
country’s history are not favourable for agronomy, and the physical and chemical
characteristics of the natural vegetation which make it unsuitable for feeding
livestock or for other forms of commercial usage other than cut flowers.
The investigation to be described is a preliminary study of one aspect of
fynbos nutrition: the nitrogen status of the family Proteaceae and its comparison
with that of non-fynbos families, in an attempt to understand this feature of the
PENNE Sn OEOt ee ee
Accepted for publication 31st October, 1977.
143
144 Journal of South African Botany
nutritional strategy of the fynbos employed in coping with the severe nutritional
limitations of the soils of their vegetational area.
In this study the following four aspects of the nitrogen nutrition of representa-
tive species of Proteaceae have been investigated:—
1. The spectrum of free nitrogen compounds found in the leaves of the species,
the quantities of these compounds present and a comparison of these data with
that of non-fynbos plants.
2. The spectrum and relative quantities of the translocatory nitrogen compounds
responsible for conveying nitrogen from soil to leaf via the xylem stream of
the root and stem, and a comparison of these data with relative data obtained
for non-fynbos representative species.
3. The effect of feeding additional supplies of nitrate or ammonium ions to the
shoots of these plants to ascertain whether these supplies can be utilised and, if
so, their influence on nitrogen metabolism in the leaf.
4. The nitrate reducing properties of the leaves of these plants and an assessment
of the role of nitrate in leaf nitrogen nutrition.
PLANT MATERIAL and METHODS
Four species of Proteaceae, Protea lepidocarpodendron L., Protea laurifolia
Thunb., Leucadendron xanthoconus O. Ktze. and Brabeium stellatifolium L. were
investigated. Protea lepidocarpodendron and Leucadendron xanthoconus were
growing at 300 m on the N.W.-facing slopes of Orange Kloof, Table Mountain, in
soil derived from Table Mountain Sandstone. Protea laurifolia and Brabeium
stellatifolium were growing respectively on the N.E.-facing slopes and the river
valley floor of Happy Valley, Bains Kloof in Table Mountain Sandstone derived
soil at an altitude of 630 m. Sampling was carried out during April and May 1976,
i.e. in late autumn.
Leaf Sampling and Amino Compound Extraction. When leaf samples were
removed from plants in the field, they were immediately killed and frozen by
immersing them in liquid nitrogen. These frozen samples were returned to the
laboratory where known weights of leaf material were homogenized in cold 80%
ethanol (100 mf 80% ethanol for 5 g leaf material) using an Ultra Turrax
homogenizer. The preparations were allowed to extract for 24 h at O ° in a sealed
flask with occasional shaking, filtered and reduced in volume to 10 m& under an
airstream.
Xylem Sap Extraction. Xylem sap was extracted from the plants by cutting off
leafy twigs in the field and applying pressure to them inside a Scholander Bomb.
The xylem sap extruded from the cut surface of the twig stem was collected and
frozen immediately.
Soil Nitrate Analysis. Soil samples were collected in plastic bags from under
the experimental plants and frozen immediately by immersing them in liquid
trogen, thus killing the soil bacteria. These were transported to the laboratory,
A study of the nitrogen nutritional status 145
dried at 30 °C and extracted by shaking in distilled water (1:1 weight/volume) for
30 min. Nitrate analysis was performed on soil extracts and xylem sap using an
Orion Nitrate Probe coupled to an Orion Model 701 pH Meter.
Amino Compound Analysis. Amino acid and amide content estimates of leaf
extracts and xylem sap were made on a Beckman 120C Amino Acid Analyser
using the lithium buffer methodology described by Lewis (1975).
Nitrate and Ammonium Ion Feeding. Nitrate ion and ammonium ion solutions
were fed to detached shoots of Leucadendron xanthoconus in the field as follows:
Shoots were detached from plants at 10h00, their severed ends recut under
water to prevent xylem airblocks and placed into beakers containing either a 200
pg N m£1KNOs solution or a 200 wg N m£~* NH,C1 solution. These shoots were
allowed to photosynthesize in sunlight alongside the plants from which they had
been detached for 8 hours, when they were frozen and extracted as described
about. Reference shoots were removed from the plants for analysis at the
beginning of the experiment and at the end of the 8-hour period.
A further experiment in which nitrate was fed continuously to detached shoots
of L. xanthoconus for 24 h was performed in a ‘‘Conviron’’ controlled environ-
ment cabinet under constant light (16 000 lux), a temperature of 16 °C and a
relative humidity of 80%.
The duplicated results of these feeding experiments are reported in Table 3.
15 N Isotope Studies on Nitrate Utilisation. In this experiment a leafy shoot of L.
xanthoconus was fed a solution of 200 wg %N mf K 'NO; for 24 h in a
controlled environment cabinet under the conditions described above, to determine
whether the shoots of this plant are able to reduce and utilise nitrate.
After extraction, the amino compounds of the extract were separated on a 150
x 1,8 cm Beckman M-84 ion-exchange resin column and collected in a fraction
collector. The amino compound samples were prepared for “N atomic emission
analysis by Kjeldahl digestion and ammonia distillation. These analyses were
performed on a Statron (Packard) atomic emission spectrophotometer following
oxidation of the ammonia with sodium hypobromite by the method of Faust
(1967).
RESULTS AND DISCUSSION
Free Amino Compounds in the leaves of P. lepidocarpodendron and L. xantho-
conus
The levels of free amino compounds occurring in these leaves are indicated in
Table 1. Also shown in this table are the relative figures for a non-fynbos plant
typified by Datura stramonium L. growing in fertilised agricultural soil near
Cape Town. The same amino-compound extraction procedure was used for Datura
as for the fynbos plants. : .
From this table it can be observed that the free amino acid and amide levels in
the leaves of P. lepidocarpodendron and L. xanthoconus are extremely low when
146 Journal of South African Botany
TABLE |
The concentrations (wmol g fr. wt. +) of the free amino compounds of the leaves of two
members of the Proteaceae compared with Datura stramonium.
Amino Acids Leucadendron Protea Datura
xanthoconus lepidocarpodendron stramonium
Aspartate 0,13 0,16 2,67
Threonine 0,10 0,07 1,16
Serine 0,30 0,26 0,94
Asparagine 0,04 0,07 0,69
Glutamate 0,30 0,36 3,68
Glutamine 0,03 0,09 1,45
Proline — — —
Glycine 0,26 0,25 0,08
Alanine 0,10 0,13 0,48
Valine 0,01 0,02 0,01
Cystine — — —
Methionine — — —
Isoleucine 0,03 0,03 0,26
Leucine 0,03 0,03 0,07
Tyrosine 0,02 0,02 0,13
Phenylalanine 0,02 0,03 0,04
Lysine 0,25 0,15 0,13
Histidine 0,10 0,05 0,04
Arginine 0,01 0,01 1,00
compared with those of D. stramonium, with serine, glycine and glutamate
dominating the amino acid spectrum.
This low free amino compound level is an indication that the level of N
metabolism activity in these fynbos plants is low in comparison with that of plants
adapted to living under more favourable soil N nutritional conditions.
Free Amino Compounds in the Xylem Sap of P. lepidocarpodendron, P. neriifolia,
L. xanthoconus and B. stellatifolium
Table 2 shows a quantitative comparison of the amino compounds, nitrate and
ammonium ions being transported from root to leaf via the xylem sap in the four
Proteaceae species with those of D. stramonium. In the Proteaceae species the
extremely low concentrations of nitrogen compounds found in the xylem sap are
immediately obvious, indicating the very restricted supply of nitrogenous nutrients
to the shoot in comparison with a plant adapted to a more favourable nutritional
environment.
In all four fynbos plants, glutamine is the dominant amino compound present
in the xylem sap, reflecting the importance of this amide as a translocator of
reduced nitrogen from root to leaf in many plants. The major utilisable nitrogen
carrier appears to be ammonium ions.
The unreduced nitrogen supply to the leaf in the form of nitrate present in the
xylem sap, is greater than the total reduced nitrogen supply (amino compounds +
147
of the nitrogen nutritional status
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Fic. 1 (a)
Nitrate content of xylem sap of Leucadendron
xanthoconus (LX) (two samples) Protea lepi-
docarpodendron (PL) Brabeium stellatifolium
(BS) Protea laurifolia (PN)
Fic. 1 (b)
Nitrate content of soils in which the plants
were growing.
Fic. 1 (c)
pH of soils in which the plants were growing.
A study of the nitrogen nutritional status 149
ammonium), but there is strong doubt as to whether this supply can be utilised by
the leaf as the experiments to be described indicate.
The quantity of nitrate present in the xylem sap appears to be directly related
to soil nitrate content and soil pH. Fig. 1 shows the correlation between xylem
nitrate content, soil nitrate content and soil pH. The lower nitrate levels in the low
pH Bains Kloof soils are probably a manifestation of the well known phenomenon
of nitrifying bacteria inhibition in acidic soils.
The Effect of Nitrate and Ammonium Ion Feeding on Nitrogen Metabolism in
Photosynthesizing shoots of L. xanthoconus
The results of the nitrate feeding experiments in which 200 wg N mf“! was
fed to photosynthesizing shoots via the xylem stream in the form of potassium
nitrate are reported in Table 3. An inspection of this table shows no significant
differences between the aminograms of experimental plants with their controls
after a feeding period in the light of 8 h. After a 24 h feeding period there is a
slight increase in the level of the N-incorporating amino acid, glutamic acid,
indicating a small degree of nitrate ion utilisation by the shoots of L. xanthoconus.
The results of the ammonium feeding experiments in which 200 wg N mf?
was fed to photosynthesizing shoots in the form of ammonium chloride are also
shown in Table 3. An inspection of this table shows a considerable increase in the
glutamine levels of the experimental plants compared with the control plants,
indicating that the additional ammonium supply can to a large extent be incorpo-
rated into shoot metabolism. Glutamine has been shown to be an important
contributor of reduced nitrogen to plant amino acid synthesis via the glutamate
synthase pathway (Lee & Miflin, 1974) and is known to accumulate in plants
receiving high nitrogen nutrition (Lewis & Berry, 1975).
Nitrate Reduction by Shoots of L. xanthoconus
The ™N enrichment of three important free amino acids of the leaf of L.
xanthoconus after feeding K’°NO3 (99 atom % excess) at the 200 wg N mf" level
for 24 h is shown in Table 4. These enrichments are extremely low for a 24 h
feeding period and confirm the poor nitrate processing properties of L. xantho-
conus shoots. It nevertheless appears that the shoots do have limited nitrate
TABLE 4
5N enrichment of glutamate, aspartate and threonine in leave
K ™NO (200 wg °N m7) through the xylem stream for 24 h.
s of L. xanthoconus fed
15N CONC.
(ATOM % EXCESS)
GLUTAMATE S
ASPARTATE i
THREONINE 1,
NOW
Journal of South African Botany
150
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A study of the nitrogen nutritional status 151
reducing properties which may be further induced after prolonged exposure to
high nitrate concentrations.
CONCLUSIONS
If the species investigated in this report can be considered representative of the
Proteaceae, it would appear that this family has adapted to the poor nutrient status
of the soils in which it grows by evolving a low intensity nitrogen metabolism to
correspond with a slow growth habit. It would also appear that the shoots of L.
xanthoconus are unable to utilise to any great degree high levels of nitrogen
supplied to them, especially if the nitrogen is provided in the form of nitrate.
Other workers (Groves & Keraitis, 1977) have shown, indeed, that high levels of
nitrogen (250 wg N mf’) and phosphorus feeding can prove fatal to Banksia
serrata, an Australian member of the Proteaceae, confirming the restricted nitrogen
processing potential of the family.
REFERENCES
Faust, H., 1967. Probenchemie '°N-markierter Stickstoffverbindungen im Mikro- bis
Nanomolbereich fiir die emissionsspektrometrische Isotopenanalyse. /sotopen-
praxis 3: 100-103.
Groves, R. H. and Keraitis, K., 1976. Survival and growth of seedlings of three
sclerophyll species at high levels of phosphorus and nitrogen. Aust. J. Bot. 24:
681-690.
Lea, P. J. and Mir.in, B. J., 1974. An alternative route for nitrogen assimilation
in higher plants. Nature, Lond. 251: 614-616.
Lewis, O. A. M., 1975. An ™N-“C study of the role of the leaf in the nitrogen
nutrition of the seed of Datura stramonium L. J. exp. Bot. 26: 361-366.
Lewis, O. A. M. and Berry, M. J., 1975. Glutamine as a major acceptor of
reduced nitrogen in leaves. Planta 125: 77-80.
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JIS. Afr. Bot. 44 (2): 153-156 (1978)
A NEW SPECIES OF PECTINARIA
P. V. BRUYNS
ABSTRACT
A new species of Pectinaria Haworth is described from the Calvinia district.
UITTREKSEL
"N NUWE PECTINARIA SPESIES
‘n Nuwe Pectinaria Haworth spesies vanaf Calvinia-distrik, word beskryf.
INTRODUCTION
Pectinaria, one of the smaller genera in the Stapelieae, is not known to occur
outside the Cape Province. Norti of the Ceres-Karoo it is represented by P.
saxatilis N.E.Br. around Vanrhynsdorp and by P. articulata Haw. var nama-
quensis N.E.Br. in the Springbok district. A few isolated records exist of P.
saxatilis from the Calvinia-Loeriesfontein area.
Pectinaria exasperata Bruyns, sp.nov.
Caules aliquando conferti aggregati vel repentes sub terram, tum surgentes
supra terram, partes sub terram albae, sine angulis manifestis, 5 ad 10 mm crassae,
partes supra terram erectae, 10 ad 15 mm crassae apud planitiem terram,
coartantes ad culminem rotundatum, armatae parvis dentibus, glabri; flores partim
vel omnino subterranei; pedicelli glabri, 2 ad 5 mm longi cum floribus; sepala
acuta, glabra; corolla 18 ad 20 mm longa; tubus ater purpureo-ruber basi, cetero
pallidus puniceus, 10 ad 12 mm longus, coartans paululum prae initio lobis,
papillatus intus cum prominentibus papillis; lobi aliquando conjuncti extremis, 7
ad 8 mm longi, 3 mm lati basi, papillati prope basim; corona lobi exteriores
minutes, bidentates; /obi interiores 2 mm longi, atri purpureo-rubri.
Type material: CAPE—3119 (Calvinia): about 20 miles north of Calvinia (—BC),
Bruyns 1345 (NBG, holo.).
Stems sometimes in a dense clump or else creeping underground from a central
group of stems for about 40 mm or longer, then emerging above surface.
Underground portion white, without obvious angles, 5-10 mm thick, above-
ground portion erect, 10-40 mm long, dull grey-green, distinctly 4-angled, 10-15
mm thick at ground level, tapering (often abruptly) to a rounded top, armed with
Accepted for publication 1st November, 1977.
153
154 Journal of South African Botany
(4)
Fic. 1.
Pectinaria exasperata:
(1) Corolla x4,5.
(2) Staminal column X18.
small sharp teeth on the angles, glabrous; flowers partially or entirely sub-
terranean, arising in groups from a subterranean peduncle up to 10 mm long;
pedicels glabrous, 2-5 mm long when with flowers, elongating to 10 mm after
fertilisation. Before seed production elongate to 20-25 mm (coming just above
soil surface) finally lengthening to 50 mm when follicles develop; sepals 3 mm
long, 1 mm broad at base, acute, glabrous; corolla 18-20 mm long; tube
cylindric-ovoid with deep purple-red base, the rest very light pink, 10-12 mm
long, 4-7 mm broad at widest, narrowing slightly to 4-6 mm just below base of
lobes, papillate within with prominent papillae covered with sub-globose, round-
topped spicules, papillae becoming smaller towards base of tube, those in
purple-red basal portion of tube each topped with a single hair up to 1 mm long;
lobes 7-8 mm long, 3 mm wide at base, with a few papillae near base, very light
pinkish-grey, with 5 intermediate decurved lobes up to 1 mm in length; outer
corona lobes minute, bifid; inner corona lobes 2 mm long with a dorsal projection
| mm in length, incumbent on anthers and meeting above stigmatic surface, dark
A new species of Pectinaria 1)3)
Pectinaria exasperata.
purple-red; follicles 100-120 mm long, with purple stripes on a light yellow
background; seeds 4 mm long, 2 mm broad, with papillate margin, upper and
lower surfaces rugose.
DISCUSSION
The structure of the inner corona lobes and the 4-angled stems place P.
exasperata in the group within the genus Pectinaria consisting of P. pillansii
N.E.Br., P. saxatilis N.E.Br., P. stayneri Bayer and P. breviloba R. A. Dyer. The
partially to completely subterranean flowers are reminiscent of P. pillansii, a
species further characterised by thick, decurved, intermediate corolla lobes. This
latter feature is present to a much lesser degree in P. exasperata, but is not
constant in that some flowers collected had very small and insignificant interme-
diate lobes. However, the erect, quadrangular above-ground portion of the stem
and papillate corolla separate it from P. pillansii as well as from P. saxatilis and
P. stayneri. .
The growth habit of P. exasperata is very similar to that of P. breviloba. Both
species creep underground with erect stem ends protruding at intervals. However,
stems of P. exasperata are much thicker. As far as the flowers are concerned these
two species differ in shape and size of the corolla and in colour. The inner corona
lobes with considerable dorsal processes are similar to those of P. saxatilis as
156 Journal of South African Botany
recorded by Bayer and Plowes (1975) from Vanrhyn’s Pass. In size they are
similar to those of P. saxatilis from Vanrhyn’s Pass as well as to those of P.
pillansii and P. breviloba.
The name P. exasperata is adopted on account of the remarkably papillate
inner surface of the corolla, visible even to the naked eye, a feature not found in
any other member of the group to which it belongs.
ACKNOWLEDGEMENTS
Many thanks are due to Mr M. B. Bayer of the Karoo Garden, Worcester for
generous help on numerous occasions. I wish also to thank Mr and Mrs P. Kotze
of Jaagvlak, Calvinia for their kindness and hospitality.
REFERENCES
Waite, A. and SLOANE, B. L., 1937. The Stapelieae. 2. Pasadena, California: Abbey San
Encino Press.
Bayer, M. B. and PLowes, D. C. H., 1975. Some Stapeliads of the Southern and
South-Western Cape. Excelsa 5: 76-80.
JIS. Afr. Bot. 44 (2): 157-185 (1978)
SOME PROBLEMS OF SOUTH AFRICAN PTERIDOPHYTA
W. B. G. JACOBSEN
ABSTRACT
A number of South African Pteridophyta are discussed with the object to point out a few
taxonomical, nomenclatural and distributional problems in the existing literature.
UITTREKSEL
SOMMIGE PROBLEME MET SUID-AFRIKAANSE PTERIDOPHYTA
*n Aantal Suid-Afrikaanse Pteridophyta word bespreek met die doel om ’n paar
taksonomiese, nomenklatoriese en verspreidingsprobleme in bestaande literatuur aan te
toon.
INTRODUCTION
The Lycopodium gnidioides complex is found to contain three different forms.
A study of original description and type as well as experimental cultivation is
called for to decide whether form 1 or form 3 have to be put on to subspecific or
varietal level or are merely ecotypes. Form 2 appears to be identifiable with L.
gnidioides L.f. var. pinifolium (Kaulf.) Pappe & Rawson.
Two separate forms of Asplenium erectum Bory ex Willd. are recognised and
described and again it is suggested to embark on a study of types and experimental
cultivation to solve the problem of eventual varietal status of one or the other.
An attempt is made to straighten out a certain amount of confusion with regard
to the Southern African species of the genus Polystichum. Six species are
discussed, of which one is described as a new species. A key to the six species is
provided to facilitate identification in the field and in herbaria. The distribution of
P. setiferum var. fuscopaleaceum sensu Schelpe is found to extend far into South
Africa. Montane forms of this species and of P. pungens sensu Sim are described.
The Dryopteris inaequalis complex is found to contain two well distinguished
forms, a large one with triangular fronds and distinct asymmetry of the basal
pinnae (the common forest form) and a small fern with elliptical fronds and
symmetrical basal pinnae. The forms have been kept separate under two specific
epithets in the past, but were combined by Schelpe (1970) under the name D.
inaequalis (Schlechtend.) Kuntze. It is suggested to separate the two forms again
and to clarify their nomenclature and status.
A form of Athyrium scandicinum with a well developed caudex from the
Eastern Transvaal is described. The unusual development of a caudex is ascribed
to climatic conditions.
Accepted for publication 10th November, 1977.
Wad
158 Journal of South African Botany
TE —
Fic. 1.
A, Lycopodium gnidioides L.f. Form 1 Note the closely adpressed, rigid leav
long, fairly thick fertile spikes. Erasmus Kop, ae Tree aa Pee cos .
Lycopodium gnidioides L.f. Form 3 Note the slightly patent leaves and the long fertile
spikes which are thinner than those of Figure A. Pirie Forest, Eastern Cape, Jacobsen 4582;
C, Lycopodium gnidioides L.f. var. pinifolium (Kaulf.) Pappe & Rawson. Pilgrims Hill,
rae Transvaal, D. Mousse 52; D, Dryopteris inaequalis (Schlechtend.) Kuntze? Note the
ick, creeping rhizome, the elliptical-lanceolate shape of the frond and the slightly reduced
symmetrical basal pinnae. Hogsback Forest, Eastern Cape, Jacobsen 4576.
Some problems of South African Pteridophyta 159
DISCUSSION
The Lycopodium gnidioides Complex
Lycopodium gnidioides is extremely variable in Southern Africa and attempts
have been made at various times to subdivide it into several species or varieties.
The author after studying his own material as well as that in the National
Herbarium, Pretoria, found that there are three rather distinct forms which are
illustrated on Fig. 1 A, B and C.
Form I coincides with Sim’s (1915) plate 177 and is characterised by its
pendant, rigid, yellow-green branches, which are often fertile to one half of
their length of 300-800 mm. The leaves are tightly adpressed. The fertile spikes
are 3-4 mm thick, the sporophylls are lanceolate, acute and keeled. A typical
representative is shown on Fig. 1, A.
Form 2 compares in habitus with Huperzia stricta (Bak.) Tard. as illustrated in
Flore de Madagascar et des Comores, 1971, Fig. IV, 5-8, and is characterised
by its erect, rigid, yellow-green branches, with fertile spikes of only 50 mm
length and 3—4 mm thickness. The maximum total height is 300 mm. The
fertile spike is often pendulous. The leaves are closely adpressed, the sporo-
phylls ovate, shortly acute and keeled. (See Fig. 1, C).
Form 3 has pendant, herbaceous, dark green branches, 400-800 mm long of
which the fertile spike may take up one third at only 2-3 mm thickness. The
leaves are loosely adpressed or somewhat patent, the sporophylls are lanceolate,
acute, not keeled and slightly reflexed at the tips (Fig. 1, B).
In view of the rather confusing evidence in the pertaining literature a more
detailed discussion of the species seems warranted. The species has been known
for a long time and its first description originates from the younger Linnaeus in his
Supplementum, p. 448 of 1781 on material collected by Sonnerat in Mauritius,
with long branches, so that it would be representative of either form 1 or 3 above.
From then on the species appears in the literature under a number of names. The
first mention of the variable habitus of the species was made by Schlechtendal
(1825-1832) writing that the plant may be either rigid and compressed or lax and
diffuse and that the colour of the dried plants may be green or yellowish-green.
His illustration (tab. 2) shows a pendant plant with fairly lax habitus and
elongated, fertile spikes. Sim (1915) also depicts on Plate 177 a fairly lax plant
with long fertile spikes and a similar illustration is brought by Tardieu-Blot (1971)
for L. gnidioides on Fig. V, 12. It appears therefore fairly well established that the
species in its original concept is generally accepted to be a fairly long, pendant
plant, with elongated fertile spikes, growing as epiphyte or chasmophyte in shady
forests.
160 Journal of South African Botany
Form 2
There is in South Africa and in the Malagassy Republic a stout, short, erect
form, with short, pendulous, quadrangular fertile spikes, which grows in full
sunlight on rocks on mountain tops or in streambeds, which although similar to L.
gnidioides in its typical form is entirely different in habitus (Fig. 1, C). A plant of
this type was described by Kaulfuss (Enum.fil. Chamisso, p. 7, 1824) as Lycopo-
dium pinifolium, which had strobili of 12-2 inches length. This species was
reduced by Pappe & Rawson in 1858 to a variety of L. gnidioides, of which they
give the following description:
‘‘Leaves densely imbricate, coriaceous, decurrent, glossy, spikes two inches
long, sessile, standing in pairs, branches dichotomous, fastigiate, curved at the
apex, bracts keeled. Stem erect, 10-12 inches high.”’
This description corresponds in every detail with the mountain top form. In
1882 Baker described a similar plant from mountains in Madagascar as Lycopo-
dium strictum and distinguishes this species from L. gnidioides as follows:
L. strictum: ‘‘Stems tufted, erect, poorly branched, 15—30 cm long, with the
leaves 6-9 mm in diameter, leaves oblong or linear oblong, obtuse,
ascending, imbricate and densely crowded, thick and rigid in texture,
6-9 mm long, midrib obscure. Spikes pendulous, single, 24-44% mm
in diameter, bracts broadly ovate, very thick, closely imbricate,
1%-2% mm long.”’
L. gnidioides: ‘‘Stems pendulous, 30-60 cm or more long, several
times dichotomously forked, with the leaves 12,5 mm in diameter.
Leaves oblong-lanceolate, moderately dense, ascending, obtuse, thick
in texture, glossy 9-12,5 mm long, midrib distinct. Sporangia in lax
spikes, which are often 15 cm long, in the axils of thick ovate-oblong
bracts, 3%—4% mm long.”’
Following the recent practice of splitting the family Lycopodiaceae into two
subfamilies Lycopodiaceae s.s. and Huperziaeceae (Rothm., Repert. Sp. Nov. 66:
236, 1962). Tardieu-Blot in Adansonia ser. 2, 10: 20 (1970) accepts Baker’s L.
strictum under the new combination Huperzia stricta (Bak.) Tard., but in 1971 (FI.
de Madagascar et des Comores, p. 30) considers Lycopodium pinifolium KIf. as
synonymous with Huperzia gnidioides (L.f.) Trev. On Fig. IV, 5-8 (p. 31) she
illustrates Huperzia stricta. The deviating habitus of the species agrees in the main
with a number of the author’s own specimens as well as with many sheets at PRE
(Fig. 1, C). There is, however, some disagreement between recorded measurements
of leaves and sporophylls between Tardieu-Blot, Schelpe (in a personal communi-
cation to Mr P. Vorster of the Botanical Research Institute, Pretoria) and the
author as the following tabulation shows:
161 Some problems of South African Pteridophyta
Tardieu-Blot Schelpe Author
Leaves:
Length mm 8-10 12-13
Width mm 2,5-3 1,75-2
Sporophylls:
Length mm 4
Width mm MHS)
Schelpe found slightly smaller dimensions in the Malagassy material than in the
South African forms, but there is a rather formidable difference between his
figures and the author’s, whose specimens originate mainly from the Eastern
Transvaal.
Leaving the general acceptance of the subdivision of the Lycopodiaceae and of
the genus Lycopodium to be settled by taxonomists and adhering for the time
being to the concept of the genus Lycopodium sensu lato the question arises
whether the form 2 should be maintained as a separate species, should be put on to
subspecific or varietal level or as Schelpe (1970) has done, should be combined
with L. gnidioides. Morphological differences between it and what is generally
considered to be L. gnidioides are slight, the forest plant usually having slightly
larger leaf dimensions. But the decisive factor appears to be the totally different
habitus, which may, of course, be due merely to an adaptation to different
ecological conditions, in which instance the plant would have to be classed as an
ecotype of L. gnidioides. Transitions to the forest forms, however, seem to be
lacking, so that the author is inclined to propose acceptance as a variety of L.
gnidioides, a view, which Schelpe seems to support—at least for the Malagassy
plant. Kaulfuss’ L. pinifolium, which was reduced by Pappe & Rawson to L.
gnidioides var. pinifolia was based on a type from South Africa. This varietal
epithet has preference before Baker’s epithet ‘‘sfrictum’’. The author proposes,
therefore, to reinstitute the name Lycopodium gnidioides L.f. var. pinifolium
(Kaulf.) Pappe & Rawson for this plant.
Further specimens of this variety in the examined herbaria are:
Author: Mount Sheba Nature Reserve, Jacobsen 4449; Ohrigstad Nature Reserve,
N. Jacobsen 4242; Elandshoogte Plantations, Jacobsen 4772.
PRE—Blauwberg, Cape, Leeman 108; Koegelberg, Cape, v.d. Merwe s.n., Table
Mountain, Cape, Esterhuyzen 15633; Port Edward, Strey 8646; Port Edward,
Umtamvuma River, Nicholson s.n.; Lusikisiki, Marais 967, Kaapsche Hoop,
Wager 85; Kaapsche Hoop, Pole-Evans s.n. ; Swaziland, Compton 26005, Pilgrims
Rest, Mousse 52; Mariepskop, van Son 1751c; Mariepskop, Killick & Strey 2389;
Mariepskop, Meeuse 9862; Mariepskop, v.d. Schijff s.n.; Chimanimani, Rhodesia,
6 000 ft, Thompson s.n.; Chimanimani, 5 600 ft, Mitchell 270; PRE 2697, sine
loc. (very typical).
162 Journal of South African Botany
Forms 1 and 3
While a separation of form 2 from L. gnidioides has also been proposed by
previous workers, the long stemmed forest form has so far always been considered
as a uniform species. As mentioned in the introduction there are, however, two
distinct forms. Which of the two represents the typical L. gnidioides? Is it the
rigid, yellow-green form 1 (Fig. 1, A) or the herbaceous form 3 (Fig. 1, B)? The
latter is in some ways similar to the illustration of L. ophioglossoides by Schelpe
(1970, Table 3) and was initially mistaken for this species by the author. Marriott
4878 at PRE was likewise misidentified as this species. Only on comparison with
other well authenticated L. ophioglossoides at PRE was the difference clearly
revealed. The tendency to misidentify the plant indicates that we have here a much
frailer form than the stiffly rigid, yellow-green form. Apart from the strong
differences in habitus, some observations and measurements show slight differ-
ences between the two forms:
Form | Form 3
Leaves:
Shape lanceolate linear-oblong
Tip acute rounded
Margin slightly inrolled not inrolled
Length mm 12-13 9-15
Width mm 1,5—2,5 2,259—2,5
Sporophylls:
Spike 3—4 mm diameter 2-3 mm diameter
Shape ovate-lanceolate broadly ovate
Tip acute acute
Length mm 4-5 2-3,25
Width mm 2 SZ
Keel present absent
As with L. gnidioides var. pinifolium the morphological differences could be
considered too slight to warrant separation and again the difference in habitus is
the decisive factor. The frailer plant cannot be considered as an ecotype as the
habitats are the same for both forms. It seems, however, that the rigid form is
confined to higher altitudes than the frailer form. A final decision whether the two
forms are merely ecotypes cannot be reached at this stage, but could possibly
result from experimental cultivation from stem tip cuttings in an atmosphere of
high humidity. Such experiments are unfortunately out of reach to the author.
A geographical separation does not seem to exist as specimens of both forms
in the author’s collection and at PRE originate from a wide range of localities,
from the forests of the south-western Cape, through the Ciskei, Transkei, Natal
into the eastern Transvaal and from there into tropical Africa.
The following localities for the two forms can be cited:
Some problems of South African Pteridophyta 163
Form 1:
Author: Erasmus Kop, Transvaal, Jacobsen 4391.
PRE—Hottentots Holland Mountains, Stokoe 1653; Nkandla, Acocks 11804: Gras-
kop, Eastern Transvaal, Strey s.n.; Erasmus Kop, Eastern Transvaal, Hardcastle
14; Boshoff Road, Hill 1629c; Haenertsburg, Sim, PRE 4331; Blaauwberg,
Transvaal, Dyer 9223; Kenya, Maas Geesteranus 5422; Lake Kivu, Congo,
Humbert 7488; sine loc., v.d. Schijff 7361.
Form 2:
Author: Pirie Forest, Eastern Cape, Jacobsen 4582; Hogsback Forest, Jacobsen
4559; 4560; Nkandhla, Jacobsen 4344; Pilgrims Rest, Jacobsen 4598.
PRE—Knysna, Bottomley s.n.; Caledon, Pillans 6342; Table Mountain, Marloth
6930; Pirie Forest, Sim 1439, 750, 665c, 666c; Port St. Johns, Mogg 4667; Port St.
Johns, McLoughlin 7656; Port Shepstone, Strey 5824; Uvongo, Strey 821; Swart
Umfolosi, Codd 1561; Natal Drakensberg, Marriott 4878; Kranskop, Strey 4243;
Pilgrims Rest, v.d. Schijff 6307; Pisanghoek, Transvaal, Bosman 3957; Vulcan
Karisimbi, Congo, Jean Louis 5433; Reunion, Schlieben 10867.
To assist field observations a key for the subdivision of the three forms is
supplied.
Stems erect, spikes occasionally
PendulousratithcstipSmeeae eer eee racer rel tele eter Form 2
L. gnidioides L.f.
var. pinifolium (Kaulf.)
Pappe & Rawson
Stems pendant ;
Leaves rigid, coriaceous, adpressed, plant drying yellow-green ......... Form |
Leaves soft, herbaceous, slightly patent, plant drying dark green ....... Form 3
Asplenium erectum Bory ex Willd.
Some confusion with regard to this species is encountered in the existing
literature. It appears as if two rather different forms are described by the same
name. Schlechtendal (1825-1832, p. 28, t.15) depicts under this name a fern with
fairly large, oblong, falcate, acute, serrate pinnae, but mentions in his description
and discussion of the species that there are two forms, one with large pinnae as
illustrated, the other with much smaller pinnae, which he interprets as merely
immature specimens as transitions exist.
Pappe & Rawson (1858) describe the species as having shortly stalked,
lanceolate, sub-falcate, acuminate, bluntly serrate pinnae and refer to Schlechten-
dal’s figure.
164 Journal of South African Botany
Since 1900 descriptions and illustrations of Asplenium erectum have been
published by Sim, p. 145, t.48 (1915), Tardieu-Blot, p. 219, Fig. XXX, 1-3
(1958) and Schelpe, p. 175, t.53, var. usambarense (1970). Sim places the species
into the much-varied Asplenium lunulatum complex as var. erectum (Bory) Sim,
while the other two authors refer to it under the name Asplenium erectum Bory
and A. erectum Bory ex Willd. respectively. While Sim’s illustration, however,
clearly resembles that of Schlechtendal, Tardieu-Blot and Schelpe show a small
fern, with small, oblong, rounded, serrate pinnae, quite different from Schlechten-
dal’s and Sim’s figures. Schelpe (1967) discusses the variety usambarense and
finds that the type material from the Mascarene Islands is intermediate between the
tropical African var. usambarense and the South African plants referred by Sim
and others to A. erectum. He agrees that a new evaluation of specific and
subspecific concepts will have to be done.
When studying collected own material, as also herbarium material at PRE and
at the University of Pretoria it was found that there are indeed two well separated
forms which in spite of many common features, such as rhizome, the green-
winged margins of the greenish-grey rhachis, descrescent pinnae a.o., show
considerable differences in total length of frond, as well as in the size and shape of
pinnae as the following tabulation demonstrates:
No. of spec. Length of Length of mid. | Width of mid.
Jacobsen lamina mm pinnae mm pinnae mm Tip Origin
3036 300 13 5) round Vumba
3898 290-350 15—17,5 6-7 round Chirinda
3124 220 16 6 round Chirinda
4353 270 12 5 round Nkandhla
4284 180-190 13-14 4-5 round Port St
Johns
2339 220 16-18 5) acute Tsitsikamma
4541 440 28-35 9 acuminate Hogsback
4533 300 23 9 acute Hogsback
4566 420 42-43 8 acuminate Hogsback
4452 400 38-40 10 acuminate Pilgr. Rest
4456 395 38 10 acuminate Pilgr. Rest
4386 590 32-35 9 acuminate Pilgr. Rest
4589 acuminate Zoutpansbg.
The difference is also clearly demonstrated on Fig 2, A and B, showing Jacobsen
4456 from the Mount Sheba Nature Reserve, Eastern Transvaal with acuminate
pinnae of up to 38 mm length and Jacobsen 4284 from Port St Johns with rounded
to sub-acute pinnae of up to 14 mm length. The longly pinnate form from
Hogsback agrees closely with Sim’s drawing, t.48
With regard to the length of the fronds Tardieu-Blot seems to have included
both forms in her description (200-600 mm), which in turn would explain her
inclusion of Sim’s illustration into the synonyms although there is a sharp contrast
I — AB
Some problems of South African Pteridophyta 165
EiGa2
A, Asplenium erectum Bory ex Willd. Large-pinna form, Pilgrims Rest District, Eastern
Transvaal, Mount Sheba Nature Reserve, Jacobsen 4456. Scale in cm; B, Asplenium
erectum Bory ex Willd. Small-pinna form, Mlolweni Forest, Port St. Johns, Jacobsen 4284.
Scale in cm.
to her own illustration. It would be interesting to know whether intermediate types
occur in Malagassy which seem to be lacking in South Africa.
A distributional separation in South Africa does not exist as both forms have
been found in the Woodbush of the eastern Transvaal and in areas as close
together as Nkandhla and Qudeni or George and Tsitsikamma.
In view of many common features it seems that both forms should be left in
the Asplenium erectum complex, but that studies of the original descriptions and
types as well as experimental cultivation should be done to reveal whether
transitions exist or whether the placing of one or the other of the two forms on to
varietal level is justified.
The Genus Polystichum Roth
The following notes should not be considered as an attempt to revise the
Southern African species of this notoriously difficult genus, but are designed to
help in the understanding and distinction of the various species in Southern Africa.
They are based on field and herbarium observations. A key to the presently
distinguishable species using simple morphological features is designed, which
166 Journal of South African Botany
includes three clear cut and well authenticated species, two species referred to as
sensu their particular authors and one new species.
In the latter instances the question of correct names may ultimately depend on
a further study of types and even then the final answer on these difficult and often
hybridising species will require cytotaxonomical studies and spore investigations
as well as possibly experimental cultivation.
It is felt, however, that pending the conclusion of such detailed studies, the
provided key will be of use to separate the various members of the genus in the
field. The species as defined are discussed with regard to their salient features and
their distribution. The pinnules of the latter three species vary much in size and
shape within fairly wide limits and some of these variations are shown on Figures
7, 8 and 9.
KEY TO SPECIES
ie Frond ipinnatepaass cece ect or seer oer eret P. macleaii (Bak.) Diels
1a Eronditwo-pinnateltosthree-pinnatiiidiener ese Sees rer
2 Rhizome, stipe and rhachis scales black ..... P. luctuosum (Kunze) Moore
2? SeRhizomeystipe andschachisiscalesibrownse ae eee eerie acer
3. Sori practically exindusiate or indusia evanescent at a very early stage
4 Pinnule margins bluntly crenate ....... P. zambesiacum Schelpe
4’ Pinnule margins sharply serrate, acute, mucronate or aristate.....
pe ult Lies ca tgeco ussianad ae Secuneucre Meter oe P. transkeiense Jacobsen
BES onibwithsmorespersistentaindUSiaseee ener
4 Sori reddish-brown, Indusia yellowish, flat, disc shaped plant
usually densely set with foxy-red scales, drying green ..........
Beer cfc ean P. setiferum var. fuscopaleaceum sensu Schelpe
4’ Sori brown, indusia light to dark brown with dark centre, funnel-
shaped, plant often set with pale brown scales, plant drying black
on upper side and silver to olive-grey on under side............
L REMC hati hence SUA Oe oe eee P. pungens sensu Sim
DISCUSSION OF SPECIES
Polystichum macleaii (Bak.) Diels in Nat. Pfl. I, 190 (1899) (wrongly spelt
Macleanii); Sim, Ferns S. Afr. 2. ed., 120, t.31 (1915).
Aspidium Macleaii sensu Sim, Ferns of S. Afr. 1. ed., 170 (1892).
This fern with its pinnate, hard, leathery fronds and sharply serrate pinnae is a
precise type and offers no problems in South Africa. Sim quotes it from the
Transkei (Bazija). The author has seen it mainly in a number of places in the
Eastern Transvaal. It has so far not been found further north and is not mentioned
in Flora Zambesiaca.
Polystichum luctuosum (Kunze) Moore, Ind. Fil.: 95 (1858); Sim, Ferns S. Afr.
2. ed. 117, t.28 (1915); Tardieu, Fl. Madag. Polypod. 1; 320, t.45, Fig. 9-12
(1958); Schelpe, Fl. Zamb. Pterid. 228, t.64, Fig. C (1970).
This is also a well defined and easily recognisable species with its characteris-
tic coat of black to dark brown rhizome, stipe and rhachis scales, the longly
Some problems of South African Pteridophyta 167
tapering apex of the frond (see Fig. 3, A and B), the often falcate pinnae and the
oblique and costa-decurrent pinnules.
Its distribution ranges from the Eastern Cape through the Transkei and Natal
into the Eastern Transvaal and into Rhodesia, where it is rare. It is also known
from Madagascar. It is always confined to deep shade in forests, often growing in
ravines or near streams and is on the whole not common.
Fic. 3.
A, Polystichum luctuosum (Kunze) Moore. Habitus, showing | the ovate-lanceolate,
attenuate fronds. Blyde River Forest Reserve, Pilgrims Rest District, Eastern Transvaal,
Jacobsen 4365. Scale in cm; B, Polystichum luctuosum (Kunze) Moore. Centre of lamina.
Note the characteristic black scales on the rhachis and the acroscopically enlarged basal
pinnules. Blyde River Forest Reserve, Pilgrims Rest District, Eastern Transvaal, Jacobsen
4356. Scale in cm.
Polystichum zambesiacum Schelpe in Bol. Soc. Brot., Ser. 2, 41: 215 (1967);
Schelpe, Fl. Zamb. Pterid.: 226, t.64, Fig. A (1970). . .
This fern, only recently described from Rhodesia, is well characterised by its
usually fairly dissected, lobate pinnules with bluntly crenate margins, which
separate this species clearly from the following three species, Aven all have
sharply serrate or aristate pinnule margins (Figs 7-9). To Schelpe s clear descrip-
tion the author would like to add some observations on dried material. All
specimens seen dried almost black on the dorsal and dull olive green on the
168 Journal of South African Botany
ventral side with slightly darker veins and a slightly chagrined surface. Indusia are
obviously evanescent at a very early stage, as they were seen only in one
specimen, where they were of half the size of the sorus.
The pinnules vary from almost undivided to pinnatifid (compare Schelpe’s
illustration in Flora Zambesiaca, Table 64, Fig. A,), are usually triangular with
rounded to acute apices, the margins bluntly serrate. The rhachis is fairly thin and
slightly flexuous with few brown scales, which also may occur on the costae (Fig.
4, A).
Initially there seems to have been some notion that this species was synony-
mous with P. ammifolium Poir. Encycle. 5: 554 (1804) (Schelpe, 1969), although
this author (1967) in his annotations to the description of the new species
maintains that P. ammifolium is apparently endemic to Mauritius. Still earlier
Alston & Schelpe (1952), however, regard P. aculeatum sensu Sim as synony-
mous with P. ammifolium. Sim’s illustration of P. aculeatum (t.26), which with its
heavy cloth of hair-like scales and its pinnules set at right angles to the costa
corresponds best with P. setiferum var. fuscopaleaceum sensu Schelpe 1970: 226,
however, seems to point to the synonymy of the latter two species. Schelpe (1970)
accordingly does not quote any synonyms and concludes that the distribution of P.
SEES aN nee Se ee,
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i ee \
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i) ae j
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2 7
er BEN Cs f
i Fix nN y
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\ S DISTRIBUTIONS OF
Ya POLYSTICHUM ZAMBESIACUM- __O
a POLYSTICHUM TRANSKEIENSE ----@
Phe a are eS |
see Fic. 10.
Distributions of Polystichum zambesiacum and Polystichum transkeiense.
Some problems of South African Pteridophyta 169
zambesiacum is confined to Rhodesia, Malawi and Mozambique. As Fig. 10
shows the species does not occur south of 20° latitude and is separated by a wide
distributional gap from the somewhat similar new species P. transkeiense as will
be discussed below.
The fern is represented in the examined collections by the following speci-
mens:
Author: Imbeza Forest, Penhalonga, Rhodesia, Jacobsen 3838, 3838a; Vumba
Mountains, Rhodesia, Jacobsen 3033, 1454, 3037; Inyanga, Rhodesia, Jacobsen
3812.
PRE—Vumba Mountains, Chase 1102; Pungwe, Schweickerdt 2412.
Whereas the previous three species are well defined and easily recognisable,
with only moderate variations in the size and shape of the pinnules, the following
three species are sometimes difficult to separate and show great variations in shape
and size of pinnules, as Figs 7—9 demonstrate. It is only by extensive studies of
herbarium material that the author can now claim to be able to distinguish between
the various species in most instances. Least affected by variations is P. setiferum
var. fuscopaleaceum sensu Schelpe which on account of its persistent bright green
colour in dried specimens and the abundance of foxy-red to light-brown scales on
rhizome, stipe, rhachis and costae is rather easily recognised. The new species P.
transkeiense usually dries green, but does turn dark on occasions. P. pungens
sensu Sim always dries black or dark olive green on the upper side and silver grey
to light olive green on the under side.
Polystichum transkeiense Jacobsen sp. nov.
Rhizoma elongata repens, paleis pallide vel atro-brunnescentis acuminatis vel
caudatis leviter dentatis obsitis. Frondes caespitosi arcuati herbacei. Stipes usque
ad 580 mm longus, viridis vel rariter pallide fuscus, glabrus vel ad basim paleis
dispersis lanceolatis acuminatis usque ad 13 mm longis obsitis. Lamina 480 mm
usque ad maxima 650 mm longa et 300 mm lata, in ambitu ovata acuta, pinnis
basalis aliquantum reductis et deflexis bi-pinnatis et leviter vel profunde tripinna-
tifida. Pinnae anguste oblongae, usque ad 240 mm longae et 60 mm latae ad
apicem longe attenuatae et caudatae. Pinnulae petiolatae in ambitu ellipticae vel
obliquo-triangulares late acutae, ad basim inaequaliter cuneatae glabrae leviter vel
profunde pinnatifidae, lobis serratis acutis vel aristatis, lobo acroscopico basale
maximo. Rhachis straminea vel viride tenua flexuosa paleis longis fimbriatis
fusci-castaneis ad basim pinnae concentratis. Sori circulares, c. 1 mm diam.,
indusium evanidum. Sporangia annulis castaneis transversis striatis et paraphysis
atris ad apicem cellulis albis translucentis praeditae.
Type: TRANSKEI, (Port St. Johns): 67 m, Jacobsen 4301 (PRE).
170 Journal of South African Botany
Also from the following localities:
Author: Mount Sheba Nature Reserve, Jacobsen 4454; Qudeni, Jacobsen 4460;
Nkandhla, Jacobsen 4355; Port St. Johns, Jacobsen 4322; Hogsback, Jacobsen
4546.
PRE—Komati Poort, Muller 2107; Qudeni, Fischer 802; Buccleuch, Natal, Sim
3641; Port Shepstone, Strey 5994; Port St. Johns, Wilkins 40; Port St. Johns,
Flanagan 2473; Port St. Johns, McLoughlin 5/36; Egossa Forest, Strey 8869;
Lusikisiki, Strey 6718; Visch River, Zeyher, coll. Sim 524c; Worcester District,
Cooper 1736; Table Mountain, Cape Town, Marloth 6936.
The fern is distinguished from P. pungens sensu Sim by its practically
exindusiate nature, the softly herbaceous character and its tendency to grow
isolated in deep shade and not in large clusters in less shaded positions as is the
case with P. pungens. From P. setiferum var. fuscopaleaceum sensu Schelpe it is
separated by the scarcity of brown scales, sometimes being practically glabrous
and its flexuous thin rhachis (Fig. 4, B and C). From P. zambesiacum Schelpe,
which the fern resembles most, it is clearly distinguished by the sharply serrate
and usually distinctly aristate or at least mucronate pinnule lobes.
The plant must have been observed by Sim (collected by him at Buccleugh,
Natal, under number 3647, PRE) as it occurs quite frequently in Natal and the
Transkei, but it is apparently included in his description of P. aculeatum, which
judging by his illustration on Plate 26 is representative of what is now known as
P. setiferum var. fuscopaleaceum sensu Schelpe.
There are distinct distributional variations, the Natal and Transkei forms
having the largest, least scaly fronds and most subdivided pinnules, while the
shape of the pinnules of the Hogsback forms tend to resemble P. pungens (Fig.
7). The few specimens from the Transvaal are distinctly more scaly and the
pinnules are only faintly lobed (Fig. 4, C). The distribution of the species is so far
somewhat disjointed, as there are a few gatherings from the southwestern Cape,
then a more or less continuous belt from the eastern Cape through the Transkei
into Natal and again a few localities in the eastern Transvaal (Fig. 10). These in
turn are separated by a wide gap from the similar P. zambesiacum in Rhodesia and
Mozambique.
Fic. 4.
A, Polystichum zambesiacum Schelpe. Centre of lamina. Note the strongly divided pinnules
with rounded lobes and the thin flexuous rhachis. Vumba Mountains, below Castle
Mountain, Rhodesia, Jacobsen 3033. Scale in cms; B, Polystichum transkeiense Jacobsen
sp. nov. Centre of lamina. Similar to A, but lobes sharply serrate-aristate. Note the thin,
glabrous, flexuous rhachis. Port St. Johns, Jacobsen 4301. Scale in cm; C, Polystichum
transkeiense Jacobsen sp. nov. Centre of lamina. Undivided pinnules of the Transvaal form,
but characteristic is the slightly scaly, thin, flexuous rhachis. Mount Sheba Nature Reserve,
Pilgrims Rest District, Eastern Transvaal, Jacobsen 4454. Scale in cm.
Some problems of South African Pteridophyta ail
2 Journal of South African Botany
3
(1-3)
ee ‘
Jacobsen 430/
Port St. Johns |
O- 6.
Al. GF
(4-6)
Jacobsen 4322
Port St. Johns Me
/0.
[L [Pp
14.
//.
CP WS. ,
a
(13-15) |
Jacobsen 4454
YUP (10-12) Mt. Sheba
Jacobsen 4355
Nkandla !
Jacobsen 4546
Hogsback
FIG. 7.
Polystichum transkeiense. Variations in size and shape of acroscopic pinnules. Note the
sharply serrate and mucronate to aristate character of the pinnule lobes and the strong
variations in size of pinnules. Natural size. LP = Lower pinnule, CP = Centre pinnule, UP
= Upper pinnule.
Some problems of South African Pteridophyta 7/3)
Polystichum setiferum var. fuscopaleaceum sensu Schelpe in Fl. Zamb. Pterid.
226, t.64, Fig. B (1970).
Polystichum aculeatum sensu Sim in Ferns S. Afr. 2.ed., 115, t.26 (1915).
The likelihood of the synonymy of P. aculeatum sensu Sim and P. setiferum
var. fuscopaleaceum has been discussed in the foregoing (see under P. zambesia-
cum). The latter name was published by Schelpe as comb. nov. et stat. nov. P.
setiferum (Forsk.) Moore ex Woynar var. fuscopaleaceum (Alston) Schelpe in
Bolm. Soc. Broteriana Ser. 2, 41: 216 (1967). To avoid ambiguity as to the true
nature of the species discussed, however, the author has preferred to refer to the
species under the above name.
The fern is fairly easily recognised by its usually heavy coat of light-brown to
foxy-red scales on rhizome, stipe, rhachises and costae, the almost rectangular
setting of the pinnules on the costae and the retention of its green colour
when dry. Variation of pinnules in size and shape is not as pronounced as in
P. transkeiense and P. pungens sensu Sim. (Fig. 8).
PRE. 7300 PRE. J.RH. Acocks. 23447
Hex Fiver Mts. (524m. Compassberg - Hanover District. 1920m.
PRE. 1867 PRE. Hafstrom & Acocks /966
Marija, Lesotho Drakensberg National Park.
Jacobsen. 4420 Jacobsen. 3087
Mount Sheba. 1/645m. Melsetter - Rhodesia. 16/4 m.
Fic. 8.
Polystichum setiferum var. fuscopaleaceum. Variations in size and shape of ened
iy - . . . °
acroscopic pinnules. Note the deviation from the usual 90 ° angle formed between costa an
basiscopic side of pinnule in the montane form from the Cape. Natural size. Abbreviations
as for Fig. 7.
According to Schelpe (1970) the distribution of the fern is confined to the
higher tropical African mountains. A thorough study of the author’s material and
of material at PRE has extended the range of the species well into the subtropical
and even temperate ranges of South Africa, the latter mostly in a slightly diverging
174
Journal of South African Botany
montane form. A comparison of A and B on Fig. 5 as well as the following
tabulation proves beyond doubt that the tropical form extends into the Eastern
Transvaal.
Jacobsen 3864
Jacobsen 4485
Locality
Altitude
Stipe
Stipe
scales
Lamina
Pinnae
Pinnules
Rhachis
Sori
Banti South, Rhodesia
1870 m
brown below, stramineous above
large, oblong-lanceolate, acuminate,
dark to light brown, 10 x 3 mm
and linear, filiform, smaller, acu-
minate, light brown
arching, herbaceous, 500 x 200
mm, oblong lanceolate-acuminate,
basal pinnae hardly reduced
100 x 23 mm, narrowly oblong,
attenuate, basal acroscopic pinnules
slightly larger
closely spaced, slightly petiolate, up
to 12 X 7 mm, cuneate at base,
forming a right angle, auriculate
acroscopically, oblong, broadly
acute, serrate-aristate, glabrous ven-
trally, softly hairy dorsally
stramineous with numerous reddish-
brown, lanceolate to filiform scales
on rhachis and costae
about 1 mm diameter, indusia fallen
Mount Sheba Nature Reserve, East-
em Transvaal
1935 m
brown below, greenish-stramineous
above
large, oblong-lanceolate, acuminate,
dark to light brown, 12 x 3 mm
and linear to filiform, smaller acu-
minate, light brown
herbaceous, 580 xX 230 mm, lan-
ceolate, longly acute, basal pinnae
hardly reduced
110 x 25 mm, narrowly oblong,
acute to attenuate, basal acroscopic
pinnules slightly larger
fairly closely spaced, up to 15 x 6
mm, cuneate at base, forming a
right angle, auriculate acroscopi-
cally, oblong, broadly acute,
sharply serrate-aristate, glabrous
ventrally, sparsely pilose to gla-
brous dorsally
stramineous, densely set with red-
dish-brown linear to hair-like
scales with laciniate outgrowths at
the base, up to 6 mm long on rha-
chis and costae
about 1 mm diameter, indusia fallen
off due to mature state off due to mature state
It is evident that Jacobsen 3864 agrees with Schelpe’s (1970) description and
is, therefore, P. setiferum var. fuscopaleaceum. With the exception of the almost
glabrous pinnules and the slightly larger frond Jacobsen 4485 (Fig. 5, B) agrees
well with No. 3864 and must likewise be considered to be the species. Other
gatherings from the Transvaal are Jacobsen 4428, a more juvenile specimen, also
from Mount Sheba Nature Reserve, where the erose indusia are well preserved,
Some problems of South African Pteridophyta 175
EIGHS!
A, Polystichum setiferum var. fuscopaleaceum sensu Schelpe. Centre of lamina. Note the
characteristic, almost rectangular setting of the lower edge of the pinnules to the secondary
costae. Banti South Endemic Forest, Umtali District, Rhodesia, Jacobsen 3864; B, Poly-
stichum setiferum var. fuscopaleaceum sensu Schelpe. Centre of lamina, showing the
typical rectangular setting of the pinnules and the strongly scaly rhachis. Mount Sheba
Nature Reserve, Pilgrims Rest District, Eastern Transvaal, Jacobsen 4485. Scale in cm.
Jacobsen 4255 from Ohrigstad, 4332 from Maviriestad near Ermelo. Jacobsen
4298 from Karkloof in Natal finally extends the range of the species into much
lower altitudes (610 m) and further south. A study of the Pretoria material showed
gatherings from Mount Hope, Upper Zwart Kei River, 1615 m (Galpin 5621) and
from Hogsback, Eastern Cape Province, probably from an altitude of 1 160 m
(Dahlstrand 2902). The range of distribution of the typical form of the species is
thus extended into the Eastern Cape Province (see Fig. 11) and into far lower
altitudes than was known so far.
The following additional localities of the species are:
Author: Melsetter, Jacobsen 3087; Banti South, Rhodesia, Jacobsen 3879; Mount
Sheba Nature Reserve, Jacobsen 4498 and 4420.
PRE—Sabi Gorge, Wager 25; Barberton, Wager 151; sine loc. PRE 746;
Maviriesstad, Polt 4848; Northern Transvaal, Repton 5B; Woodbush, Reynolds,
176 Journal of South African Botany
DISTRIBUTIONS OF:
POLYSTICHUM SETIFERUM
VAR. FUSCOPALEACEUM
AS ABOVE. DRAKENSBERG FORM. ©
POLYSTICHUM PUNGENS
Fic. 11.
Distributions of Polystichum setiferum var. fuscopaleaceum, its montane form and of
Polystichum pungens.
PRE /0246; Pilgrims Rest, Collins 895c; Woodbush, Jenkins 919c; Sabie, Wager
48; Duiwelskloof, Scheepers 419; Natal, PRE 2/68; Zwartkop, Natal, Sim, PRE
388; Drakensberg, Esterhuyzen 26045; Lion’s River, Natal, Moll 829; Cathedral
Peak, Killick 1134 and 981 (identified by the collector as P. setiferum (Forsk.)
Woynar); Natal, Wood, coll. Sim 520c; Van Reenen, Schlechter 42c; Lidgetton,
Natal, Roberts 87f; Underberg, Natal, MacClean 684; Tugela Gorge, Hutchinson
41; Lesotho, Dieterlen 695 and 1309; Pirie, East. Province, Sim PRE 3642;
Compassberg, Acocks 23447.
While the typical form in most instances loses its indusia at a relatively early
stage, although relics may often be preserved, a montane and southern form of the
species is characterised by the retention of the indusia well into the mature state.
The author refers to this form as the Drakensberg Form of the species. Doidge
1061 at PRE shows clearly that the initial stage of the development of the sori
commences with the typical flat, disc-like, colourless, erose indusia of the typical
form of P. setiferum var. fuscoplaeaceum. The indusia later become mushroom
Some problems of South African Pteridophyta 177
shaped or slightly funnel-shaped, changing their colour from pale yellow to yellow
and reddish-brown, at which stage the likewise reddish-brown sporangia protrude
around the margins. Finally the indusia become drawn into the sori which often
have tufty white paraphyses. When the sporangia have fallen off an elongated
black scar is left along the veins of the pinnules.
The Drakensberg Form is even more densely set with scales than the typical
form, having a heavy coat of ovate-lanceolate light-brown to foxy-red scales on
thizome and lower stipe, 12 x 5 mm in size, with darker brown centre stripe and
translucent lighter brown edges. The scales on the rhachis are filiform, foxy red,
up to 6 mm long. The frond is narrowly lanceolate, with closely set acuminate
pinnae and overlapping pinnules, set at 60 ° to 90 ° to the costae, but mostly still
revealing the characteristic 90 ° angle in places.
The persistence of the indusia, the dense cloth of scales and the closely set
pinnae and pinnules point to an adaptation to colder climatic conditions, but may
also be caused by growth in more exposed, less shady habitats which prevail in the
range of distribution of the Drakensberg Form at altitudes from about | 700 m in
the southwestern and southern Cape to more than 2 760 m in the Drakensberg.
Specimens have been collected in an area extending from the Hex River Moun-
tains in the southwestern Cape towards the Compassberg and the high portions of
the northeastern Cape Province, the Orange Free State, Lesotho and Natal and
northwards as far as Wakkerstroom in the southeastern Transvaal (see Fig. 11).
The following gatherings have been made:
PRE—Oshoek, Wakkerstroom District, Devenish 2538 and 195; Thaba N’chu
Mountain, O.F.S., Roberts 2998; Clarence, O.F.S., van Hoepen 18230; Mt. Aux
Sources, Bottomley, PRE 2497; Goodoo, Mt. Aux Sources, Doidge 1061;
Drakensberg National Park, Hafstroem & Acocks 1966; Cathedral Peak, Drakens-
berg, Killick 981 (identified by the collector as P. setiferum (Forsk.) Woynar);
Giant’s Castle, Snyman 1281; Cathedral Peak, Rush 2030 and 2300; Gudu Forest,
Natal National Park, Edwards 325; Lesotho, Schmitz 402 (identified by the
collector as P. setiferum (Forsk.) Woynar); Lesotho, Guillarmod 690; Lesotho,
Bevis 102; Molteno, Flanagan 1681; Top of Mt. Koudveld, Herb. Marlothianum
PRE 2/70; Highbury, Polela District, Natal, Doidge PRE 9; Barkley East, Galpin
6934: Matatiele, Acocks 12207; Queenstown, Hangklip Mountain, Roberts 2012;
Compassberg, Coetzee PRE 11456; Montague Pass, Schweickerdt PRE 4705; Hex
River Mountains, Esterhuysen 14885.
Polystichum pungens sensu Sim in Ferns S. Afr. 2. ed.: 116, t.27 (1915);
Aspidium aculeatum Sim in Kaffr. Ferns 46 (pro parte) (1891); Aspidium acule
atum var. pungens sensu Sim in Ferns S. Afr. 1. ed.: 166 (1892). .
This fern is generally referred to in the South African literature (for instance
Schelpe, 1952) and herbaria as P. /ucidum (Burm.) Becherer. The type of this
178 Journal of South African Botany
(1-3)
Jacobsen 2337 Knysna.
&
(4-6)
Jacobsen 4535 Hogsback.
ff 7
“one neo
ee
ne 4
(7-9)
Jacobsen 4557 Hogsback.
Tor Doone.
‘p i
(10) (11-12)
PRE. 387 Zwaartkop Natal. PRE. Zwaartkop Pletermaritzburg Oistrict.
USs /4. 15.
(13) (14 - 15)
PRE. 9313 Woodbush. Tv/.
PRE. 746/ Woodbush. Tv/.
Fic. 9.
Polystichum pungens. Variations in size and shape of acroscopic pinnules. Note the strongly
deviating Natal and Tor Doone forms. Natural size. Abbreviations as for Fig. 7.
Some problems of South African Pteridophyta 179
plant, named Asplenium lucidum by Burmann fil. is in Geneva and Becherer gave
only a very brief description of it. Aspidium pungens Kaulf. was based on a
specimen collected by Chamisso in the Cape and is now in Leningrad. As the
author is not able to examine either of the types and to avoid confusion as to the
plant referred to, it is here quoted as Polystichum pungens sensu Sim.
In his description Sim mentions that this fern has given rise to much confusion
in the past due to its extreme variability, especially in the size and shape of its
pinnules of which Fig. 9 tries to convey an impression. This could possibly lead to
the subdivision into a number of forms or even varieties. The author, however, is
of the opinion that the varying characteristics are on the whole impersistent in
distribution and nature, with the exception of a montane form which will be
discussed below.
One of the safest ways of distinguishing P. pungens from P. setiferum var.
fuscopaleaceum and P. transkeiense in the dried state is the colour, which with
P. pungens invariably turns blackish or dark olive-green on the upper side and
silvery- to olive-grey on the underside. The indusia are practically persistent,
peltate, stalked, funnel-shaped, initially colourless to light brown, later dark
brown, often with a dark centre, only drawn into the sori at a late stage. The
sporangia are brown with a light to dark brown annulus.
The pinnules are strongly varying in size, 13-30 mm long, 6-10 mm wide at
the base, their shape varying from non-lobate forms (Natal and Katberg) to
strongly subdivided lobate forms (Tor Doone, Hogsback area) as Fig. 9 demon-
strates. The typical and most common form of the Cape (Fig. 6, A) has hard,
leathery laminae with sharply aculeate pinnule lobes and a heavy coat of
yellow-brown to reddish-brown scales on rhizome and lower stipe, which extends
to the full length in young fronds. The less typical forms of Natal and Transvaal
are not as leathery and more herbaceous.
P. pungens occupies on the whole slightly lower altitudes in the Cape and
Natal than P. setiferum var. fuscopaleaceum. In the Transvaal the fern becomes
much less common and is found sporadically in the forests of the eastern
Transvaal as far north as the Woodbush. P. pungens is the most common fern in
the southern Cape and the forests of Kaffraria and assumes in this respect the role
which further north is taken up by Dryopteris inaequalis. Here it usually forms
large clusters, often fully exposed to sunlight above the forests and on top of
mountains amongst stones.
The typical form is represented by a great number of specimens:
Author: Mount Sheba, Transvaal, Jacobsen 4433; Hogsback, Jacobsen 4535,
4553, 4557 (Tor Doone form); Storms River, Jacobsen 2331; Knysna, Jacobsen
2337; George, Jacobsen 2314, VOD, VENT, FIND
180
Fic. 6.
A, Polystichum pungens sensu Sim. Right centre pinnae. Note the oblong, falcate, sharply
serrate-aristate pinnules. Woodville Forest Reserve, Wilderness, Cape, Jacobsen 2314.
Scale in cm; B, Athyrium scandicinum (Willd.) Presl. Caudex, 15 cm high. Mount Sheba
Nature Reserve, Pilgrims Rest District, Eastern Transvaal.
Some problems of South African Pteridophyta 181
PRE—TRANSVAAL: De Hoek Forestry Reserve, Schweickerdt PRE 4689:
Woodbush, De Hoek, Schweickerdt 1852c; Woodbush, Wager PRE 7461; Wood-
bush, Jenkins 919.
NATAL: Ngoya, Zululand, Medley-Wood 10886 (pinnae undivided, hardly aris-
tate); Zwartkop, Sim PRE 3560, 387, 11026, 9259; Drakensberg, Bottomley PRE
5018; Pietermaritzburg, Strey 8429 (Tor Doone form); Maritzburg, Sanderson,
coll. Sim 530c (Tor Doone form); Eshowe, Laura PRE 6421; Winterskloof,
Doidge PRE 54 (Katberg form).
TRANSKEI AND CAPE: Port St. Johns, Wager PRE 2996; Engcobo, Tembuland,
McLoughlin 1045c; Kentani, Pegler 270 (pinnules rather broad, sharply serrate-
aristate); Komgha, Flanagan 290; Hogsdack, Dahlstrand 1550, 1853; Pirie, Sim
PRE 3554; Pirie, Sim 533c, 515c (with foxy-red rhizome, stipe and rhachis
scales); Pirie, Sim 5/6c (very broad pinnules, but distinctly aculeate and drying
black on upper side); Mount Kemp, Pirie, Sim 528c; Pirie, Sim PRE 3559; Dohne
Hill, Sim 512c, 513c; Keiskammahoek, Wells 3343; Amabele District, Hardcastle
293 (small form, with broad, poorly divided pinnules); Frankfort, Sim 512c;
Katberg, Young, coll. Sim 518c (with broadly rhomboidal, rectangular based,
poorly divided, but strongly aristate pinnules); Zuurberg, Rogers 1478c; Albany
District, Cooper PRE 1161, 1415; Great Winterhoek, Uitenhage, Esterhuysen
PRE 10847; Storms River, Dahlstrand 1693; Storms River, Schlechter 5963;
Knysna, Marloth 5711; Knysna, Rex 840c, 838c (Tor Doone form, with dense
foxy-red scales on rhachis); Plettenberg Bay, Rodin 1191; George, Schlechter 525;
Wilderness, Mogg 11636; Ladysmith, Esterhuysen 14013; Hofmansbosch, Britten
PRE 4241; Cape Town, Marloth 541 (very large pinnules, sharply serrate,
aristate); Table Mountain, Esterhuysen 15355 (Tor Doone form).
Similar to P. setiferum var. fuscopaleaceum, P. pungens has developed a high
altitude montane form which may be distinguished from the typical form by the
almost total lack of scales on rhachis, costae and costulae. The lower part of the
stipe bears a cloth of dark reddish-brown linear to lanceolate scales. The texture of
the lamina is thick and leathery, drying in contrast to the typical form to a fairly
even matt olive-green on both sides. The pinnules may be aculeate, but are more
often only coarsely serrate without the usual bristle points of the common form.
Representative specimens are:
PRE—Tabanhlope Mountain, Natal, Wisch 184; Tugela Valley, Hafstroem &
Acocks 1970; Mont Aux Sources, Schweickerdt 760; Cathedral Peak, Esterhuysen
10199a; Cathedral Peak, Edwards 851; Rockeries Pass, Drakensberg, Edwards
2145; Golden Gate Highlands Park, O.F.S., Liebenberg 7498.
The Dryopteris inaequalis Complex.
Schelpe (1970, p. 221) describes Dryopteris inaequalis (Schlechtendal) Kuntze
as a fairly large fern with creeping rhizome, large pale brown rhizome scales, a
182 Journal of South African Botany
long stipe, a large, broadly to narrowly ovate, triangular, two-pinnatifid lamina
with the basal pinnae hardly or not reduced. While this description fits the most
common forest fern in Southern Africa, the author observed and collected a
similar, but much smaller fern in various forest localities which differs from D.
inaequalis sensu Schelpe in its present concept by its non-triangular, broadly
lanceolate frond and its distinctly reduced basal pinnae. While the large fern is
usually 3- to 4-pinnatifid (not always 2-pinnatifid as Schelpe’s description of the
Rhodesian form confusingly seems to convey), the small fern is never more than
2-pinnate to 3-pinnatifid. Its characteristics seem to agree well with the description
of Dryopteris inaequalis sensu Sim (1915, p. 106, t.18); while the characteristics
of Schelpe’s D. inaequalis are coinciding with D. elongata sensu Sim (1915, p.
104, t.7). The history of the synonymy of the two species seems to contain a
certain amount of confusion until 1964, as Table 1 demonstrates.
It is obvious that Schelpe in his later publications, combined the two species in
one which he considered to be D. inaequalis (Schlechtend.) Kuntze, thus dis-
regarding that this name referred to the small and relatively rare species of Sim
(1915) and that D. elongata sensu Sim, corresponded with the common large
forest fern. He must have come to the conclusion that the two species were not
sufficiently distinct to be separated. Sim admits that there may be confusion
between the two species when young. This was also observed by Burtt-Davy
(1926) and the author. But in all instances where adult material is present, the
distinction is easy. The author collected the small species in the Wilderness and
the Hogsback and Pirie Forests. It is clearly distinct from the tall, triangular fronds
of the common large fern by its very pronounced creeping and unusually thick (for
such small fern) rhizome, its lanceolate fronds with widely separated pinnae, of
which the basal pair is always smaller than the one above (Fig. 1, D), thus
contrasting sharply with the distinctly triangular fronds of the larger fern, where
the lowermost pinnae are always the largest. The author also had no difficulty in
separating the relatively few specimens of the smaller fern from the several
hundred specimens of the D. inaequalis complex at PRE, provided the entire frond
and possibly the rhizome are preserved.
The following specimens were recognised to belong to this type:
Author: Hogsback, Jacobsen 4576, 4571; Pirie, Jacobsen 4584; George, Jacob-
sen 2318.
PRE—Mooi River, Natal, Mogg 2160; Zwartkop, Natal, Sim PRE 3624; Frankfort,
Cape, Sim 480c; Pirie, Sim 469c; Wilderness, Mogg 11775; Keiskammahoek,
Kotsokoane 246; Woodvale Forest, Queenstown District, Galpin 8202 (determined
by Schelpe as D. inaequalis (Schl.) O. Ktze. and thought to be intermediate with
D. elongata); Ravine, 9% miles up Kowie River, Bosman PRE 2968; sine loc.,
PRE 2295; sine loc., Wells 3339; Kookfontein, Burger 961c.
183
Some problems of South African Pteridophyta
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184 Journal of South African Botany
It appears therefore, as if the re-institution of the previous subdivision into two
different types may be justified, and that a clarification of the nomenclature and
status of the two forms is called for.
The distribution of the smaller fern is quite interesting as it seems to be
confined to a relatively low altitude belt, extending from the Wilderness over the
forests of the Eastern Cape Province to Natal. Sim, in addition, quotes it from the
Swellendam area, which appears feasible, but also from the Northern Transvaal
(Woodbush) and Rhodesia as well as Mozambique. There are, however, no
records from these areas at PRE.The altitude seems confined to the span from sea
level to 1 220 m.. The habitat requirements are deep shade, either in herbless
ravines and gullies or, as frequently in the Hogsback and Pirie Forests, under a
dense herb layer and then extremely difficult to find. The plant may therefore not
be as rare as stated, but merely overlooked. The large form is common on forest
floors throughout Southern Africa, but is also found in fairly open ground, at
forest edges or on granite hills in Rhodesia, in which instances the pinnule lobes
tend to become narrower and the pinnae are slightly wider spaced.
Athyrium scandicinum (Willd.) Presl
Schelpe, Flora Zambesiaca, 1970, uses as one of the distinguishing features
between the genera Athyrium and Diplazium the absence, respectively presence of
a caudex. Although in most instances applicable it was found that Athyrium
scandicinum may develop a distinct caudex (Fig. 6, B). The author saw this species
in the Mount Sheba Nature Reserve, Pilgrims Rest District, in a streambed on the
so-called ‘‘Sheba Trail’’, shortly before this emerges from the forest into open
grassland. About four or five specimens are growing here in a cluster on large
boulders of quartzite, having caudices from 50 to 150 mm height from which large
tufted fronds of about 600 mm lengih and 230 mm width emerge. The caudices
are between 30 and 40 mm thick. The thick erect rhizome or caudex and the
merely slightly incised pinnule lobes identify the species as A. scandicinum, as the
similar A. schimperi has a creeping rhizome and strongly and deeply incised
pinnule lobes.
References to this unusual habitus could not be found in the corresponding
literature. Sim (1915) speaks merely of a suberect paleaceous crown. Tardieu-Blot
(1958) quotes an erect rhizome and tufted fronds. More precise statements are
found in Schelpe (1970): ‘“‘Rhizome up to 5 mm in diameter, erect, with tufted
fronds and oblong etc. rhizome scales’’. Yet in 1956 Schelpe collected his No.
6002 (PRE) at Entabeni in the Zoutpansberg with the annotation ‘‘Rhizome erect
and massive’. This is apparently a similar tall specimen with fronds of 650 mm
length and approximately 300 mm width. From the use of the expression
‘massive’ the author concludes that this may well have been a plant with a small
caudex. Schelpe’s key in Flora Zambesiaca, therefore, may thus be restricted to
tropical Africa (Rhodesia, Malawi, Mozambique), but loses its validity further
Some problems of South African Pteridophyta 185
south. This geographical restriction seems to point to the possibility that the
development of a caudex by A. scandicinum may be due to climatic factors and
may take place only in the cool to cold montane forests of the Transvaal and
possibly further south? The species extends in its distribution to Natal, but here
usually at lower altitudes. Sim (1915) reports it also from Grahamstown and from
Lesotho. Specimens from the latter locality might be interesting, but are unfortunately
not represented at PRE.
ACKNOWLEDGEMENTS
The author wishes to express his sincere appreciation and thanks for the
valuable assistance received from the Director of the Botanical Research Institute,
Pretoria, Dr. B. de Winter, as well as from Mr. P. Vorster, Botanist in charge of
the Cryptogam Section of the same institute.
Thanks are also due to Prof. P. J. Robbertse of the University of Pretoria for
permission to study herbarium material at the Department of Botany.
Finally, the author would like to thank the Directors of Nature Conservation in
the Transvaal, Natal and the Cape Province for the permission to collect Pterido-
phyta at various periods through the last years and especially Mr. N. Jacobsen of
the Department of Nature Conservation of the Transvaal who during his work in
various Nature Reserves collected many interesting specimens for the author’s use.
REFERENCES
Atston, A. H. G. and ScHevpe, E. A. C. L. E., 1952. An annotated check-list of the
Pteridophyta of Southern Africa. J/ S. Afr. Bot. 18: 153-176.
BAKER, J. G., 1887. Handbook of the fern allies. London.
Burtt-Davy, J., 1926. A manual of the flowering plants and ferns of the
Transvaal with Swaziland, South Africa. London.
Pappe, L. and Rawson, R. W., 1858. Synopsis Filicum Africae Australis. Cape
Town.
ScHELPE, E. A. C. L. E,, 1967. New taxa of Pteridophyta from South East
Tropical Africa. Bolm Soc. broteriana Ser. 2, 41; 203-217.
ScHELPE, E. A. C. L. E., 1969. A revised check-list of the Pteridophyta of
Southern Africa. JI S. Afr. Bot. 35 (2): 127-140.
ScHELPE, E. A. C. L. E., 1970. Pteridophyta. In: A. W. Exell and E. Launert,
Flora Zambesiaca. London.
SCHLECHTENDAL, F. L. de, 1825-1832. Adumbratio Filicum in Promontorio
Bonae Spei Provenientum. Berlin.
Sim, T. R., 1915. The Ferns of South Africa. 2nd ed. Cambridge.
TARDIEU-BLoT, M. L., 1958. Polypodiacées (sensu lato). In: H. Humbert,
de Madagascar et des Comores. Paris.
TARDIEU-BLOT, M. L., 1971. Lycopodiacées, Huperziacées. In: J. F. Leroy,
Flore de Madagascar et des Comores. Paris.
Flore
JIS. Afr. Bot. 44 (2): 187-200 (1978)
“DRIFT SEEDS”? AND *“‘DEAR DIRTY DUBLIN’: CORRESPONDENCE
BETWEEN JOHN MUIR AND MATILDA C. KNOWLES, 1930-1933
E. CHARLES NELSON
(National Botanic Gardens, Glasnevin, Dublin, Ireland)
ABSTRACT
Between 1930 and 1933, Dr John Muir of Riversdale, corresponded with Miss M. C.
Knowles, botanist in the National Museum of Ireland, Dublin, about drift seeds. Their
letters, published herein, reveal some of the problems faced by Muir in his research on alien
beach drift, and the extent of the assistance he received from Dublin.
UITTREKSEL
“DRIFT SEEDS”’ EN *“‘DEAR LCIRTY DUBLIN’’: BRIEFWISSELING TUSSEN JOHN
MUIR EN MATILDA C. KNOWLES, 1930-1933
Tussen 1930 en 1933 het Dr John Muir van Riversdal briewe met Mej. M. C. Knowles,
plantkundige aan die Nasionale Museum van Jerland, Dublin, gewissel oor dryfsaad. Hulle
briewe onthul sommige van die probleme waarmee Muir te doen gekry het in sy studie van
uitheemse opdrifsels en die hulp wat hy van Dublin ontvang het.
INTRODUCTION
One of the most important contributions to research on long-range dispersal of
plant propagules by ocean currents was made by Dr John Muir, who carried out a
very detailed and extensive survey of alien fruits and seeds in the beach drift of
southern Africa.
John Muir (1874-1947) was born in Scotland and qualified as a doctor of
medicine at the University of Edinburgh in 1896. In the same year he emigrated to
South Africa where he practised medicine for nearly thirty years. Muir eventually
settled in Riversdale, and after retiring from medical practice in 1923, he
concentrated on botanical research. As a medical practitioner, Muir worked on
haemophilia, and according to one of his letters to Miss Knowles (see below), he
spent two months at the Coombe Hospital in Dublin in 1910.
Muir’s botanical research was wide-ranging, and gained him honorary docto-
rates from the Universities of Edinburgh and Stellenbosch. However, his interests
also included sea-shells, ornithology, genealogy, folk-lore and he collected the
vernacular names of plants, shells, and birds in the Riversdale region (see
Hopkins, 1972; van Blerk, 1972).
In their recently published work on tropical drift fruits and seeds, Gunn &
Dennis (1976) said that without Muir’s studies the beach drift of Africa ““would be
Accepted for publication 10th November, 1977.
187
188 Journal of South African Botany
virtually unknown’’. Muir visited beaches on the Mozambique, Congo and
Cameroon coasts, as well as those in South Africa (Gunn & Dennis, 1976). He
made extensive collections of ‘‘drift seeds’’, which are preserved in the University
of Stellenbosch (STE-U), and he obtained seeds from living plants to aid
identification of the beach collections. Muir published his data in a series of
scientific papers and popular articles (e.g. Muir, 1929a, 1929b, 1931a, 1931b, 1932,
1934, 1937).
CORRESPONDENCE WITH DUBLIN
A major problem faced by research workers studying “‘drift seeds’ is the
identification of specimens, which is intensified as the “‘seeds’’ are not attached to
plants whose characters could aid identification. The disseminules of genera of
Leguminosae especially are difficult to identify to species level (Gunn & Dennis,
1976). Muir’s correspondence with Dublin arose from his need for assistance with
the identification of his specimens.
It seems, from extant letters, that Muir first wrote to Ireland in order to get a
copy of a paper published by Nathaniel Colgan (1919) on Irish “‘drift seeds’’. This
first letter probably was addressed to Dr. R. LI. Praeger, the leading Irish botanist
at that time, and Praeger sent a copy of the paper to Muir; this letter is not extant.
The earliest surviving letter, written by Muir to Praeger, thanks Praeger for the
Colgan paper, and requests assistance with identification.
In 1915, Dr H. B. Guppy who was studying tropical ‘‘drift seeds’’ (see Guppy,
1917) presented to the National Museum of Ireland a collection of seeds and fruits
of West Indian plants whose propagules were recorded from beach drift in the
British Isles as well as ‘‘seeds and fruits. . . that have not yet been recorded from
the beaches of the United Kingdom though it is likely that some of them have
been picked up but have not been identified’’'. Guppy hoped that this collection
‘‘though not a large collection. . . (would) be of a little service in directing once
again the attention of botanists in Ireland to this subject’’”. Guppy (1917) noted that
he had presented this collection to Dublin, as did Colgan (1919).
Muir’s letter requesting identification assistance was passed to Miss Knowles,
the “‘Lady Assistant’? in charge of the museum’s botanical collections. Miss
Knowles studied the specimens sent in by Muir, and sent him a set of notes on the
material after she had compared the seeds with those in Guppy’s collection. In his
reply thanking Miss Knowles for her aid, Muir complained that Kew had not
given any help, even though there was a very large collection of ‘‘drift seeds’’ in
the Royal Botanic Gardens. Undoubtedly, Muir was very grateful for the assistance
given by Miss Knowles, and the friendly though sporadic correspondence that
continued until 1933 clearly indicates the rapport that arose between Dublin and
Riversdale.
In gratitude for Miss Knowles’ help, Muir sent to Dublin a small collection of
seeds and fruit, both beach drift and fresh specimens. These specimens are
“Drift Seeds’’ and ‘‘Dear Dirty Dublin’’: A correspondence 189
preserved in the herbarium of the National Botanic Gardens, Glasnevin (DBN)
along with Muir’s letter and some rough drafts of Miss Knowles’ replies; the
botanical collections of the National Museum were transferred to the National
Botanic Gardens in 1970. Muir also sent offprints of some of his papers and
articles; these are also held in Glasnevin. As noted, Muir’s own collection of
“drift seeds’’ is preserved in the University of Stellenbosch (STE-U), along with
letters from Muir’s correspondents, including Miss Knowles; no letters from Dr
Praeger are preserved in Stellenbosch (Jordaan, pers. comm.), though perhaps
none was written.
Miss Knowles’ last letter to Muir, thanking him for an offprint of one of his
papers (Muir, 1932) was written a few weeks before she died after a short illness
(Praeger, 1949).
THE LETTERS
The letters are published in chronological order, and are printed in full; some
of the rough drafts included in the Glasnevin archives aye not included. Copies of
Miss Knowles’ letters (originals in STE-U) have been placed in the Glasnevin
archives, and copies of Muir’s letters (originals in DBN) are available in the
University of Stellenbosch.
This first letter was written to Dr Praeger; it was passed by Praeger to Miss
Knowles and therefore is preserved with this correspondence.
The original spelling and punctuation of the letters are retained in this paper.
‘Leeuwarden’
Riversdale
2 June 1930
Dr John Muir -
(M.B., D.Sc.)
Dear Mr Lloyd Praeger, ; 4 :
Very many thanks for the valuable paper by Mr Nathaniel Colgan” on the larger alien
Irish Beach Drift. I am only sorry that I cannot consult Dr Guppy who passed away in 1926.
I have been unable to get a single seed named, although the Buitenzorg people in Java sent
me a nice lot of cultivated seeds and fruits.
I have just found that Barringtonia speciosa* reaches here.
I am venturing to send you a few seeds and would be very grateful if you would have
them compared with Guppy’s which he gave to the National Irish collection. If you will
return them to me I wd. send some to you later if they wd. be acceptable. hal
352:83° Kew named these near Entada sp. but they are Afzelia bijuga [= Intsia bijuga|
I feel sure. They grow when sown and the leaves are not quite the same as Intsia bijuga
from Madagascar, or the seeds are larger than those I have seen from Calcutta and the
Phillipines. Still I am nearly certain. , ae Ae 9
171.40. These must be Dioclea sp. Did Guppy give you anything like these?
255.373. These I think must be Strongylodon ruber (lucidas)? Do they resemble any
form of Mucuna urens known to you? Is the cosmopolitan polyzoon on 255 Membrani-
190 Journal of South African Botany
pora? No one in S. Africa knows it. If so Mr Colgan has solved one of my difficulties for
me. They may be Mucuna sp.
36. Must be Dioclea reflexa: black form
324. Must be Dioclea reflexa: brown form
445, 443, 26, 314: various types of Mucuna spp unknown to me and everyone I have
consulted.
Entada scandens® and Caesalpinia Bonducella (crista)’ as being abundant and well
known I have not sent nor the usual Rhizophoraceae and other mangroves.
All were picked up here. If you care to lecture on them I have not the slighest objection,
senders rights protected.
I know all the seeds mentioned by Mr Colgan, as I have Sacoglottis from Trinidad.
With Kind regards, I remain
Yours sincerely
John Muir.
They must all come from Madagascar, Malaya or E. Indies. None are African.
A note, in pen, is written on the first page and reads “*Seeds returned & notes on
comparisons with those in our collection. July 4. 1930 MCK”’.
Ard-Mhusaeum Na h-Eireann
(National Museum of Ireland)
Sraid Chill Dara
(Kildare Street)
Baile Atha Cliath
(Dublin)
July 4 1930
Dear Dr Muir,
As Dr Lloyd Praeger was leaving Dublin for some weeks when your letter & box of
drift seeds arrived he passed these on to me and asked me to make the comparison with the
drift seeds in our collections and let you know the result. As Dr Praeger has probably told
you our collection is a small one composed almost entirely of the seeds that reach our Irish
shores. These have however all been vouched for by either Mr Colgan or Dr Guppy, so
although our collection is a very limited one it is reliable so far as it goes.
It has been very interesting to compare your seeds with ours but I am afraid you will
find the results very disappointing. I have made a few notes on a separate sheet [see below]. It
will be a satisfaction to you to find that your Membranipora on No. 255 is M.tuberculata. |
sent the seed to Mr Farran of the Department of Fisheries and enclose his note about it
which you need not trouble to return’. I also showed it to Mr Nicholls who used to be
keeper of the Natural History Collections here & who specialized in this group and he too
considers it ‘‘almost certainly’? Membranipora tuberculata. 1 am indeed very sorry and
disappointed that our collection has not proved more helpful in solving your difficulties.
The seeds of these Leguminous climbers seem to vary a good deal even in the same species
which makes the identification troublesome & uncertain.
Dr Guppy was a great loss. I wonder where his collection went to?® I should have
expected Kew to get it, but it may have gone to the Natural History Museum, South
Kensington, London. Mr Ramsbottom is the present Keeper of the Botanical Collections
ag
“Drift Seeds’’ and ‘‘Dear Dirty Dublin’: A correspondence 19]
there & would I am sure help you in any way he could if you wrote to him. Dr Rendle
retired a few years ago.
I am returning your seeds as requested & hope they will reach you safely.
With best wishes
Yours sincerely
Matilda C. Knowles
Assist. in charge of the
Botanical Collections
P.S. It is very kind of you to offer us some seeds & we should be most grateful for any you
can spare us.
A rough draft of the notes on Muir’s seeds in Miss Knowles’ writing is preserved
in DBN; it is printed here. Corrections to the notes have been edited into their
correct places in the text.
Drift seeds—South African—from Dr Muir
353.83. We have nothing at all like these in our drift collection. I have also looked up
Afzelia in our Herbarium as sometimes there are seed samples attached, but our only
species is A.Africana. A.bijuga probably belongs to the section with no arillies.
171.40 We have one mottled seed similar to these but rather larger & thicker in tone,
with a darker raphe. Guppy labelled this Dioclea reflexa. Your seeds agree very well with
his description of the seeds of this species in ‘‘Plants, Seeds and Currents’ p. 131 and no.
40 though slightly smaller is similar in shape to fig. 6. in the plate at the beginning of this
book.
255, 273 These are not exactly like any seeds of Mucuna urens in our collection, they
are smaller & the raphe does not encircle the seed so much as in our samples of Mucuna but
more than in Strongylodon or Dioclea according to the descr. in Genera Plantarum of these
genera. They are rather like two seeds amongst those Guppy gave us labelled ‘*Dioclea
(probably) guianensis’’ except for being a little less round (but they seem to me to be too
round for Dioclea‘’). I do not know Strongylodon but the hilum is described in Genera
Plantarum as being semicircata while these are more than %. Mr Farran of the Fisheries
Department thinks the polyzoon encrusting No. 255 is Membranipora tuberculata.
36,324 are very like Dioclea reflexa. They are rather flatter than our specimens and 36
is more oblong in shape otherwise similar. There is one seed that is identical in every
respect with your number 324 (and one or two other very similar) amongst a sample of drift
seeds we have from Keeling or Cocos Island"', given us many years ago by the then
Director of Kew Gardens and labelled ‘‘Mucuna sp.’’ There are a number of undoubted
Mucuna seeds in the sample but those similar to your 36 are more like Dioclea reflexa |
think.
443 We have nothing like’
26 We have nothing like this
S14 ks a Es 2 i
445 more like Dioclea than Mucuna | should say”
192 Journal of South African Botany
Miss Knowles returned Muir’s seeds to him and received the following letter.
29 July 1930
Dear Miss Knowles, ine
Your letter of July 4th is just about the most helpful communication I have ever
received during two years work at the alien and local Beach Drift. Mr N. E. Brown, a very
old friend, wrote by the same mail and I hear from him nearly every week. He says that the
collection of beach drift at Kew is immense but that no one ever works at it.
Kew did not name a single one of my species and were not one tenth as helpful as you
with your small collections.
Please thank Mr Farran for naming the Membranipora.
This is just a short note of thanks and to give you a piece of what to me is important
news.
My 352 & 83 which you write about as follows ““We have nothing at all like these in
our collections’? and named at Kew ‘‘near Entada’’ is Afzelia bijuga A. Gray (= Intsia
bijuga) | received seeds from the French authorities at Madagascar ten days after I wrote to
you. You will see much about them in Guppy’s ‘‘Nar. Pacific’ II’*. They have not before
been found in the beach drift of other tropic regions (ib. 174). The Riversdale beach is
temperate and my seeds which are washed up come almost entirely from Madagascar. I
have pleasure in presenting you with one of each.
I agree with you entirely that my 36 and 324 (the latter identical you say with one of
Guppys from the Cocos Islands named at Kew Mucuna sp.) is Dioclea & not Mucuna. I cut
them open & they have no intercotyledonary cavity. The kernal floats independently of the
testa for some time, & they belong to the Sect II of the group of non adaptive seeds (Nat.
Pac. II 107). Do you remember which one in your collection is= my 324 (unspotted, dark
brown)?
I am also sending you a short paper on the Folk Lore of these seeds’’. Ent. scandens &
Caesalp. Bonducella grow in Port. East Africa. I have a pod of the former 37” long, and the
prickly pod of the latter.
I have been working through Schimpers ‘‘Die Indo-Malayisch Strandflora’’’®, a rare
book now a days which took me a year to get. Guppys paper in Journ. of Vict. Instit. in the
Keelings has so far eluded me“’. I was glad to get Colgan’s fine paper.
It is fine to see your name in Guppys P.S. and C., & to receive a letter from some one
who knew Guppy which I never did’.
In 1910 I had two months at the Coombe Hosp. Dublin & spent one of the happiest
times of my life in that unsavoury although in other respects delightful surroundings. My
recollections of Ireland have always been of the pleasantest.
Some of my seeds have gone to Madagascar, Mauritius & Calcutta.
Calcutta seeds of Afzelia bijuga & one lent me by Kew from Luzon agree; but they are
much less massive that the Madagascar ones. Either more than one species is included
under that name, & the genus requires revision, or it varies enormously.
I will send duplicates of seeds later, if Calcutta or Antananarivo’? recognise them.
With best wishes
I remain, Yrs sincerely
John Muir
Dublin. August 27 1930
Dear Dr Muir
Iam just going off for my belated summer holiday but I must send you a few lines
before I go to thank you for your very kind letter of July 29th & for the valuable additions
to our Drift Seed Collection which arrived safely a few days ago.
“Drift Seeds’’ and “‘Dear Dirty Dublin’: A correspondence 193
I am delighted to get the two seeds of Afzelia bijuga, one from Madagascar & one from
the Riversdale coast & also the two samples of Scaveola (sic)*® & I think it is most awfully
good of you to have rewarded my small efforts so generously. I shall incorporate them in
our Drift Seed Collection on my return”? & I am sure some worker at the subject will one
day bless you. Accurately named specimens are not easy to get.
I wish Guppy had lived to know that you have become so interested in this fascinating
but difficult & tedious question of the Drift Seeds. It is such a help & stimulation to have a
fellow enthusiast to discuss ones problems with. I did not know him personally—only
through correspondence.
I shall pass on your message to Mr Farran on the first opportunity I am sure he will be
glad to know he has been of use.
Dr Praeger looked in here yesterday & was asking for you. He is just back from the
Botanical Congress at Cambridge where they had a very busy week over the revision of the
Rules of Nomenclature, which they seem to have settled satisfactorily. He was much
pleased to learn that Colgan’s paper had proved of interest to you. I have promised him the
loan of your paper on the Folk Lore of Entada & Caesalpinia™.
You will have to come over & work out the Drift seeds at Kew yourself & if so you will
probably run across and renew your acquaintance with *‘dear dirty Dublin’’ & pay us a visit
here. I should enjoy going through our collection with you.
With renewed thanks & apologies for this scrappy letter. I am yours sincerely
Matilda C Knowles”®
P.S. I made a note of the seed I thought like your no 324”
Riversdale 19.8.1930
Dear Miss Knowles
Guppy’s paper ‘Dispersal of Plants as illustrated by the Flora of Keeling Atoll.”’ Journ.
of Vict. Inst. Lond. 1889 (1890) is unobtainable in S.Afr. and I have not yet got it from
Edinburgh or London. Is it very long? If not cd. you have a precis made by one of your
staff in exchange for seeds? I can send you
Canavalia obtusifolia
Ipomoea pes-caprae (fresh and from drift)
A pod of Caesalpinia Bonducella with its two seeds in situ
Strongylodon lucidus (= ruber), named since I received your letter at Calcutta, confirm-
ing my suspicions. They are the seeds of mine that you saw (see Guppy Nat. Pac. Il).
Please note for your private information that my seed 324, which you say= one of
Guppy from the Keeling Atoll in your possession, named Mucuna, but which I said is
Dioclea, is now named by the Calcutta people Dioclea javanica Bth”. My Afzelia bijuga
seeds are also now confirmed as such by them, which I had anticipated.
D.javanica is not mentioned by Guppy in any of his works, being probably under the 3
or 4 ‘‘Mucuna spp.’’ mentioned from Keeling in his paper.
I remain
Yrs sincerely
John Muir
Did I send Dr Lloyd Praeger a copy of my
paper (Bot. Survey of S-Afr.) ““Veget of
Riversdale’? He is well known by reputation
in S.Africa
194 Journal of South African Botany
Copies of Muir’s earlier papers on drift and his ‘‘Vegetation of Riversdale’’ are
preserved in Glasnevin; the copies are signed by Muir. In the archives at
Glasnevin there is a rough, incomplete draft of a reply to the above letter. This
draft is dated October 7 1930, but there is no letter with this date in the archives at
the University of Stellenbosch. It must be assumed that Miss Knowles did not
send the letter, and that Muir’s next letter reminded her about replying.
Riversdale, 19 Nov. 1930
Dear Miss Knowles
I send you the little book*® under separate cover, also in this letter a few seeds of
Ipomoea pes caprae (=I biloba) from the beach drift of Pondoland (Urnzinivuba River at
St. John’s) The bare ones are the typical: the others are fresh.
I do not know what to do about the paper of Guppy’s. It is the first of those mentioned
under his name in Plants Seeds & Currents p.vii (Dispersal . . . Keeling Island) The year
was probably 1890. I have seen the 1889 number but although there was some thing about
Keeling Island by him in it it did not deal with the drift at all.
The Vict. Instit. is more a religious society, and I cannot get a copy of the paper in any
scientific library.
Would the incumbent of St Patricks Cathedral [Dublin] not have it in their library?
With kind regards, & hoping the seeds will escape the Post Office stamp.
Yours sincerely
John Muir
Dublin, Decr 9 1930
Dear Dr Muir
Your letter with the nice little batch of seeds of Ipomoea pes Caprae and your fine
paper on “‘The Vegetation of the Riversdale Area’’ all arrived this morning. Thank you
very much indeed for them I am looking forward to reading your paper with much interest.
I was just about to write to you when your letter came to tell you that through Dr
Praeger I was able to get a short loan of Guppy’s paper on the ‘‘Dispersal of Plants as II.
by the Flora of Keeling or Cocos Islands’? and made a few extracts for you just to let you
know roughly the contents of it. These are so rough I am almost ashamed to send them as I
have not had time to type them out; but they will let you see the plan of the paper. I was
talking to Dr Praeger about how to have an abstract made for you, but he is of opinion that
no abstract is really reliable & that it would be much more satisfactory for you to see the
original paper and he said that he was quite sure if you know any member of the Linneaen
Society that he could borrow it for you. Dr Praeger says the Linnean Society constantly
lends books to specialists who are living abroad who have not a good library near. I do
hope you will be able to get the paper as it is very annoying not to be able to see the whole
literature on the subject one is studying. I am so sorry I have been able to do so little to help
you in this matter.
With kind regards and very good wishes for Christmas & the New Year
I am yours sincerely
Matilda C Knowles
“Drift Seeds’’ and ‘Dear Dirty Dublin’’: A correspondence 95
Riversdale, 3
Dear Miss Knowles, wersdale, 30 XII 1930
Many Thanks for the great trouble you have taken in making a precis of Guppy’s
article. I have ordered the book itself from Edinburgh as they once sent me the previous vol
of the series. Guppy’s own reference to his paper gives the wrong year! Schimper gives it
as ‘‘about 1890’. Thus I cd never tell them exactly what I wanted.
I am (notwithstanding the above address) in my summer abode on the Indian Ocean at
Still Bay, 25 m from Riversdale. I will send you some fruits and seeds when I return home
about Jany. 24th.
In May I hope to go to Lourenzo Marques. Did I send you seeds of Ipomoea
(Calonyction) Bona Nox, which float well but are said by Guppy to be ‘‘variable’’? I think I
did.
Would you care for the seedlings of the mangroves (Rhizoph. mucronata and Bruguiera
gymn.”° ) for the museum sheets or for the students?
Entada scandens and Caesalpinia Bonducella 1 will see in any quantity growing. I have
any quantity of seeds from the drift as well.
I have lately settled the problem for S.Afr. of Aleurites, Sophora tomentosa and
Erythrina indica.
I have had a bad set back with French red tape which will amuse you. The Director of
Agric. Mauritius (who died in Noy.) sent some of my seeds to the Chief of Agric.
Madagascar. Both are or were Frenchmen. The Postal authorities of Madagascar wd not
allow the entry of my seeds as they are contrary to regulations (fear of diseases &c). and
returned them to Mauritius. The Chief of Agric Madagascar was quite willing to examine
them but cd. not get them through the customs.
My seeds all belong to Ewarts macrobiotic group and may have been 5 to 100 years old
for all anyone knows.
The Madagscar man is the only authority living who cd. name the seeds some of which
I gave or lent you. How to manage I do not know, but neither Guppy nor Schimper ever got
their Mucuna spp. named as the genus is very difficult to determine from seeds.
Otherwise my work is finished.
The Gvt. has ordered seeds for me from Zanzibar, Ceylon, Singapore and Buitenzorg,
but the Miss Knowles’ of this world are few and far between, and it is rare to find
governments or people who will do as much for you, as you would do for them.
With very best wishes
I remain yrs very sincerely
John Muir.
Riversdale, 10.2.1931
Dear Miss Knowles,
I am sending you herewith the following from Portuguese East Africa, although they are
I am sure (being also W. Indian) all represented in your noble collection; but of interest
maybe because of their new locality: °°
6 fresh seeds of Entada scandens taken from the shrub
5 segments of the hugh pod of the same which is over 3 ft long. If you glue them
together on a flat stick they make a nice specimen. There are often 11-13 segs.
Seeds of Caesalpinia Bonducella
Pod of Caesalpinia Bonducella } y
2 fruits of Anacardium occidentale L. (Cashew Nut) This had poor powers of flotation,
but is discussed by Guppy.
With Kind regards
I remain Yours sincerely
John Muir
196 Journal of South African Botany
I have put in a pod of Canavalia obtusifolia. These fresh materials are instructive when
placed as controls against drift stuff. Of the latter I have abundance—about 600 seeds of all
kinds. Is there anything you would like?
I wish I were back at the Coombe [in Dublin], walking through the dirty streets, with
the Irish women all shouting ‘‘Good Luck’’! I will never forget Dublin, hence also these
seeds. Now I am too old alas!
Dublin March 9 1931
Dear Dr Muir
Thank you very much indeed both on my account and on behalf of the Museum for the
very nice box of seeds that arrived safely a few days ago. Especially for the pod of
Caesalpinia Bonducella (of which we had no sample) and also for the pod of Canavalia
obtusifolia.
It is an excellent suggestion of yours to mount the fresh seeds and place them as
controls against the Drift Seeds and I am carrying it out. Our collection will soon be a
‘““noble’’ one indeed thanks to the many additions you have so kindly sent us.
We are also very pleased with the sections of the Entada pod you have given us. We
already had a number of sections of a slightly parger pod, but I think yours is really better
for exhibition purposes as our show cases are not very large so I am mounting them in the
way you describe.
Dublin sends you greetings! At present it is in a whirl of excitement, which I am not
very sure is good for it, about the Hospital Sweep Stake*". I hope your old friend the
Coombe may benefit. With Kindest regards and wishing you the best of good luck
I am
yours sincerely
Matilda C. Knowles
I hope you succeeded in getting Guppys paper?
Riversdale 20.4. 1931
Dear Miss Knowles,
I am sending your hypocotyls of two Eastern Mangroves, which you may not have, see
Guppy Nat. Pac. I.
They will look well glued on cartridge paper some with fruits attached others separate:
[here there is a pen sketch to illustrate the mounting technique suggested]
You will find Ceriops illustrated in Schimper’s ‘‘Indo-malayische Strandflore’’
ae ak Rhizophora in the Western Mangrove is a different species R. margle (Guppy
0.C.)
I am going up I hope to Portuguese E. Afr. in June, when there is no malaria to have a
hunt round and see what I can get lying on the tropic beaches.
I have not succeeded in getting Guppy’s Keeling Is paper yet, but I am doing my best to
get the volume. I got the old Volume of the Journ. of the Linnean Society with Baron’s
Madagascar paper from Prof. Thoday of Bangor as a present last week.
Guppy published his papers in a religious transactions, which is perhaps the reason why
botanists do not have them! Anyhow they are almost unobtainable today I tried more than a
year to get Schimpers “Yndo-mal Strandflore’’ and at last got it in Leipzig through the
intermediations of a Hollander.
“Drift Seeds’’ and ‘‘Dear Dirty Dublin’: A correspondence 197
The papers issued by the Buitenzorg Gardens in French & German are hard to get: I
mean the Krakatoa ones.
With kind regards, I remain
Yours sincerely
John Muir
Dublin 16 Decr. 1931
Dear Dr Muir
I must apologise for not having sent you a line or two sooner to thank you for all the
interesting matter about Drift seeds that I have received from you in the last few months.
You are most kind & we are veiy grateful to you. I have received ‘‘The Romance of a
Floating Seed’’** most interesting reading. How pleased you and Mrs Muir must have been
to run your bean to earth, and what an ingenious plan you adopted to locate it. I
congratulate you on your thoroughly satisfactory piece of Research & I am glad to know
that Mucuna quadrilata is the same as Mucuna gigantea. The crocodiles and the hippopo-
tami were interesting side shows that I could have done without. You speak feelingly of the
suffering of those who have the misfortune to get the hairs of the pod into their clothes. No
wonder the natives call them “‘i-mamba’’. I must have some extracts made from your
account for my Drift seed case.
With very many thanks and the best of good wishes for 1932 from
Matilda C Knowles
For over a year, there was no correspondence between Muir and Miss Knowles.
We may presume that he had exhausted the usefulness of the very small collection
of seeds in Dublin, and, as he himself had said, his work was ‘‘finished’’.
Muir’s next letter to Dublin enclosed a copy of his paper on drift seeds (Muir,
1932), which was the result of his research, and was probably the reason for
stating that his work had finished. Muir did not end his work at that point,
however, as he published further articles including his most important paper on
drift seeds several years later (see Muir, 1934, 1937).
Riversdale 13 Febr. 1933
Dear Miss Knowles
I send herewith a paper containing two plates of seeds.
Have you by any chance a seed of Pangium edule (Flacourtaceae) collected by Guppy
in the Dutch E. Indies. aa
If you could let me see one I would send it back by the very first post, together with a
fine fruit of Heritiera littoralia for your collection <n RT
John Muir
198 Journal of South African Botany
Dublin March 31 1933
Dear Dr Muir, ; : d
Thank you very much for the paper on “‘The Beach Drift of South Africa’’ with the two
plates of seeds. These will be most useful here and helpful to all those studying Beach
Drift.
I am very sorry we have no specimens of Pangium edule. in our collections or you
should have had the loan of it with much pleasure.
I am resigning my post here on the Ist of June next and going up to Ballycastle in the
North of Ireland to live with a sister who has a little house and a nice garden there. I am
looking forward to a good rest during the summer.
If I find any interesting Drift seeds on the coast up there I shall let you know.
You have been so generous in giving us drift seeds for our collections and it is a great
satisfaction to me to know that there is such a representative set here. Thank you very much
for numerous contributions.
With Kind regards
Yours sincerely
Matilda C Knowles.
Three weeks after writing this letter Miss Knowles died, and her sister died three
days later (Praeger, 1949).
CONCLUSION
Although he stated that his work was finished in 1930, Muir did not in fact
conclude his researches then. From other unpublished letters (from H. N. Ridley to
J. Muir, in STE-U) we learn that Muir visited London and Kew at least once.
Further, his main publications on drift fruits and seeds were published between
1932 and 1937.
The co-operation indicated by the letters published here, resulted in the
assembly in Dublin (DBN) of a small collection of drift seeds and herbarium
specimens of coastal plants. Muir acknowledged the help given by the Dublin
botanists Dr. Praeger and Miss Knowles in his publications. There is little doubt
that Muir greatly valued the help given by Dublin, and the short correspondence
probably would have continued for many years had Miss Knowles not died.
ACKNOWLEDGMENTS
I am very grateful to Prof P. G. Jordaan for sending me copies of Miss
Knowles letters and for his assistance with the identification of Muir’s seeds. My
thanks are also due to Dr C. R. Gunn for his help.
NOTES
1 DBN mss. H. B. Guppy to C. W. Steele, National Museum, Dublin, 10 May
1915.
2 DBN mss. H. B. G. to The Director, National Museum, Dublin, 25 May
1915.
3
4
)
“Drift Seeds’ and “‘Dear Dirty Dublin’’: A correspondence 199
Proc. Royal Irish Academy, 35B: 29-54. 1919.
correctly Barringtonia asiatica (L.) Kurz; see Gunn & Dennis (1976).
the specimens listed have been examined by Dr C. R. Gunn and identified as
follows:
352 and 83—Intsia bijuga (Colebr.) O. Kuntze
171 and 40—Dioclea reflexa Hook.f.
255 and 373—Mucuna myriaptera Baker
36—Dioclea reflexa
324—Dioclea reflexa
445—Canavalia sp. (Muir named this Canavalia bonariensis Lindl.)
26, 314 and 443—Mucuna gigantea DC.
correctly Entada phaseoloides (L.) Merrill; for discussion of Entada spp. see
Gunn & Dennis (1976).
correctly Caesalpinia bonduc (L.) Roxb.
STE-U mss. G. P. Farran to M. C. Knowles, undated (copy only in DBN).
there are “‘drift seeds’’ collected by Guppy at Kew, as well as those already
noted in DBN. There are no Guppy specimens in BM.
in transcribing this rough draft pencil insertions have been placed into the
main text in the appropriate place. The section in brackets appears to belong here.
i.e. Cocos (Keeling) Islands in the Indian Ocean north of Australia.
Miss Knowles originally wrote “‘like Mucuna but raphe very narrow’.
deleted in rough draft.
Observations of a Naturalist in the Pacific between 1896 and 1899; vol. II
Plant-Dispersal. 1906, London.
Folk-lore of Some South African Sea Beans (Muir, 1929a): a signed copy was
sent by Muir to Miss Knowles on 29 July 1930, although Muir dated it
n2997 3" “(DBN):
Schimper A. F. W., Indo-malayische Strandflora. 1891, Jena.
Dispersal of plants as illustrated by the flora of the Keeling or Cocos Islands.
Journ. Trans. Victoria Inst., 24: 267-306 (1890).
Miss Knowles corresponded with Guppy (see Guppy, 1917) but no letters are
extant. They never met.
Tananarive, Madagascar. ,
Scaevola plumieri Vahl; Muir sent fruits from shrubs with the dried pericarp
iniact (Riversdale Coast, July 1930) and “‘stones’’ from beach drift which had
lost their pericarp (Riversdale Coast, July 1930); DBN.
material was incorporated and is still intact in DBN.
see 15 supra. Fay
a rough draft of this letter, differing slightly from the copy sent 1s in DBN.
this postscript is added to the letter on the top of the first page. It seems that
Miss Knowles wrote this letter before she received the letter sent by Muir on
19 August 1930, and that the footnote was in reply to that letter.
200 Journal of South African Botany
25 see 5 supra.
26 Vegetation of the Riversdale Area (Muir, 1929b), copy signed by Muir in
DBN.
27 Praeger worked in the National Library in Dublin, he was Chief Librarian.
28 there are no seeds of this in DBN.
29 Rhizophora mucronulata Lam., Bruguiera gymnorrhiza Lam.
30 see 5 supra; specimens in DBN.
31 a lottery run to provide funds for Dublin hospitals and prize money for
horse-racing.
32 Muir, 1931b.
REFERENCES
CoLGAN, N., 1919. On the Occurrence of Tropical Drift Seeds on the Irish Atlantic Coasts.
Proc. R. Ir. Acad. 35B: 29-54.
Gunn, C. R. and DENNIS, J. V., 1976. World Guide to Tropical Drift Seeds and
Fruits. New York: Quadrangle/New York Times Book Co.
Guppy, H. B., 1917. Plants, Seeds and Currents in the West Indies and the
Azores. London: Williams & Norgate.
Hopkins, H. C., 1972. John Muir. In: Dictionary of South African Biography.
Vol. 2; pp. 500-501. Cape Town: Tafelberg-Uitgewers Ltd.
Muir, J., 1929a. Folk-lore of South African Sea Beans. J. med. Ass. S. Afr. 3:
Murr, J., 1929b. Vegetation of the Riversdale Area. Mem. bot. Surv. S. Afr. 13.
Muir, J., 193la. Alien and Indigenous Fruits and Seeds in the South African
Beach Drift. Riversdale: private publication.
Muir, J., 1931b. The Romance of a floating seed (Mucuna gigantea). South-
western Echo (21 November 1931).
Muir, J., 1932. The beach drift of South Africa. J. bot. Soc. S. Afr. 18: 5-10.
Muir, J., 1934. The correlation of arrival sites of alien seeds with ocean currents.
S. Afr. geogr. J. 17: 13-18.
Muir, J., 1937. The Seed-Drift of South Africa and some influences of ocean
currents on the strand vegetation. Mem. bot. Surv. S. Afr. 16.
PRAEGER, R.LI., 1949. Some Irish Naturalists. Dundalk: Tempest.
VAN BLERK, N., 1972. John Muir. In: The Standard Encyclopaedia for Southern
Africa, Vol. 7, pp. 633-634. Cape Town: Nasou Ltd.
JIS. Afr. Bot. 44 (2): 201-202 (1978)
Book REVIEWS
POLLINATION MECHANISMS, REPRODUCTION AND PLANT BEEDING, by R. Frankel
and E. Galun with pp. xi+281,77 figures and 39 tables. Berlin-Heidelberg—New
York, Springer-Verlag, 1977. U.S. $26,40.
Pollination Mechanisms, Reproduction and Plant Breeding is a cloth bound volume and
the second in the series ‘““Monographs in Theoretical and Applied Genetics’’.
The authors set out to “‘furnish under one cover an integrated botanical, genetical and
breeding-methodological treatment of the reproductive biology of spermatophytes—mainly
angiosperms.’’ The book is intended for “biology and agricultural students at the graduate
level, to plant breeders and other people interested in acquiring a broader knowledge of the
reproductive biology of higher plants.”’
The first of the three chapters deals briefly with the significance of different pollination
mechanisms and more extensively with the mode of reproduction in higher plants and the
ecology and dynamics of pollination. The second chapter is devoted to self-fertilizing crops.
The evolutionary aspects and mechanism of autogamy is discussed, and the chapter is
concluded with a very apt description of emasculation and controlled pollination.
The third chapter, which comprises about two-thirds of the text, provides an excellent
detailed discussion on sexual reproduction in higher plants, control and modification of sex,
incompatibility and male sterility and the utilization of these in plant breeding.
The book is a comprehensive and up to date reference text. Nearly two-thirds of the 948
references are dated 1960 or later. There are few typographical errors and the editing
appears to have been careful. The book provides factual information which had not
previously been obtainable in one volume. The authors employ a very affective system of
headings and subheadings which enable the reader to identify specific topics without delay.
In the South African context this book should be compulsory reading for final year plant
breeding undergraduate students, post graduate students, research workers, and university
teachers specializing in this field. But priced at U.S. $26,40 it will most probably only be
libraries of research institutes and universities which will acquire copies of this publication.
H. A. VAN NIEKERK
PROCEEDINGS OF THE SECOND NATIONAL WEEDS CONFERENCE OF SOUTH AFRICA,
1977, pp. 344, with 82 figures and tables, thread sewn with paper cover, R15,00
Obtainable from the publisher: A. A. Balkema, P.O. Box 3117, Cape Town.
Conferences come and go and result inevitably in: “‘the Proceedings of .. .”.
Organising committees often seek through their choice of papers, to endow their conference
proceedings with a larger function and status—that are seldom achieved. The Pr peeeaies
of the Second National Weeds Conference of South Africa however, has a function and has
attained a status that was not really aspired to by the organisers. These proceedings set out
to be nothing more than the written record of 26 research contributions relating to a ee
of weed problems. But because of a growing realisation of the importance of weeds in
South Africa and the dearth of information about them, these proceedings find ENS
elevated almost automatically to the status of a “national review aus ee ae
reading” for anyone interested in our weed problems. They must be pee oe vee
Judged as a ‘‘national review’’ of current weed problems and researc ne ox pr oe
ings succeed in providing a fairly balanced and comprehensive coverage of the country
201
202 Journal of South African Botany
weed problems, the plants and conditions that cause them, and of new developments in
control and utilisation research. The emphasis is on the very grave threat that invading weed
species pose to our rich and beautiful Cape Flora. In view of this emphasis, it is refreshing
to also find included a paper championing one plant, black wattle, that is widely regarded as
a weed by conservationists.
Other contributions of note, include works on the chemical control of: bush encroach-
ment, Nassella tussock, wild oats; and of the weeds of vineyards, maize, wheat, clover
pastures and sugar cane. There are four papers relating to the biological control of Opuntia
aurantiaca (jointed cactus), and other pest plants. And botanical studies which provide an
understanding basic to the control of weeds, have also been included, e.g. a study of the
taxonomy of Lantana, and one on the growth kinetics of Eichornia crassipes (the water
hyacinth).
However, the coverage is not complete, and falls short of what one would expect of a
national review in that it lacks any indication of empirically evaluated research and control
priorities. And, whilst the human population explosion (the most important single factor
guaranteeing more and bigger weed problems in the future) figures in several papers,
insufficient attention is paid to the likely affect of future land use on weed problems. The
publication is also the poorer for having neither an index nor an overall summary.
Whilst these proceedings may fall short of what is required of a “national review’’ there
is no doubt at all that they must be regarded as prescribed reading for weed scientists as
well as for farmers, and conservationists who are interested in the weed problems that
threaten the environment and productivity of our sub-continent. Together with the proceed-
ings of the First National Weeds Conference of South Africa, 1974 (which is unpublished
and now almost unobtainable) they provide the first modern picture of weed problems and
weed research accents in South Africa.
With these proceedings, it is now possible to combine and extrapolate the estimates of
various authors to derive an estimate of what weeds cost South Africa annually. However
dangerous an exercise like this may be, it is important to scientist and layman alike, that it
can be indulged in.
In view of their importance, is it too late to appeal for the publication of the proceedings
of the First National Weeds Conference to supplement the work reviewed here?
The preparation of the various contributions was supervised by the individual authors,
and since printing was by offset litho minor typing errors and inconsistencies in format have
been perpetuated. These do not detract from the value of this workmanlike production. But
the cover may prove to be a little on the light side for the attention that its contents will
receive, and the use that will be made of them.
Dr. D. P. Annecke of the Plant Protection Research Institute (Department of Agricultu-
ral Technical Services), chairman of the organising committee, and Messrs. Agricura who
made a generous donation towards printing costs, are to be congratulated on this timely
publication.
M. J. WELLS
SUPPLEMENT TO AQUATIC PLANTS OF AUSTRALIA, by Helen I. Aston.
Melbourne University Press, 1973 and reprint 1977.
Reviewed: JI S. Afr. Bot. 41 (4): 273 (1975).
An eight page, loose-insert supplement applicable to both the original 1973 printing and
the 1977 reprint of Aquatic Plants of Australia has been prepared. It contains updated
information on Australian aquatic plant taxa, together with a bibliography, and is
automatically included with each copy of the reprint. Holders of the 1973 printing may
obtain a copy of the supplement free of charge from Melbourne University Press, P.O. Box
278, Carlton South, Victoria, Australia 3053.
Self-addressed envelope (10 inches x 7 inches) MUST accompany application.
JIS. Afr. Bot 44 (3): 203-279 (1978)
THE GENUS GALIUM L. (RUBIACEAE) IN SOUTHERN AFRICA
Cu. PUFF*
(Institute of Botany, University of Vienna, Austria)
ABSTRACT
A new classification of the genus Galium in Southern Africa is proposed, based on
morphological investigations, karyology, pollen measurements, and ecological observations.
Twenty taxa belonging to fourteen species are recognized: (1) G. thunbergianum, with var.
thunbergianum and var. hirsutum in East Rhodesia, Transvaal, Swaziland, Natal, Orange
Free State, Lesotho, and Eastern Cape Province; (2) G. capense, with ssp. capense in the
South West Cape and Eastern Cape Province, ssp. namaquense (stat. nov.) in the Western
Cape Province, and ssp. garipense (stat. nov.) with var. garipense and var. wittbergense
(stat. nov.) in Transvaal, Natal, Orange Free State, Lesotho, and Eastern Cape Province; (3)
G. monticolum, a rare species from the Western Cape Province; (4) G. bredasdorpense (sp.
nov.) endemic to the limestone formations in the Bredasdorp district; (5) G. amatymbicum,
a rare species from the Eastern Cape Province; (6) G. bussei, with var. bussei and var.
glabrum in Rhodesia; (7) G. scabrelloides (sp. nov.) in the South Transvaal, Swaziland,
Natal, Orange Free State, Lesotho, and Eastern Cape Province; (8) G. scabrellum in East
Rhodesia; (9) G. subvillosum, with var. subvillosum and var. subglabrum (var. nov.), (10)
G. mucroniferum, with var. mucroniferum and var. dregeanum (stat. nov.), and (11) G.
undulatum (nom. nov.) in the South West Cape; (12) G. tomentosum in the Cape Province
and South South West Africa (and Orange Free State); (13) G. chloroionanthum in East
Rhodesia; (14) G. spurium ssp. africanum, a rare taxon occurring in Transvaal, Natal, and
Cape Province.
New chromosome numbers are reported for G. scabrelloides and G. capense ssp.
garipense var. wittbergense (n = 11, 2n = 22), and G. mucroniferum vat. mucroniferum
and G. subvillosum var. subvillosum and var. subglabrum (n = 22, 2n = 44). Previously
published chromosome numbers of G. thunbergianum var. thunbergianum and var.
hirsutum and G. capense ssp. garipense var. garipense (n = 11, 2n = 22) and G.
tomentosum (n = 22, 2n = 44) are confirmed by additional new counts.
UITTREKSEL
DIE GENUS GALIUM L. (RUBIACEAE) IN SUIDELIKE AFRIKA
’n Nuwe indeling van die genus Galium in Suidelike Afrika word voorgestel, dit word
baseer op morfologiese ondersoeke, kariologie, stuifmeel afmetings en ekologiese waar-
nemings. Twintig taksa behorende tot veertien soorte word erken: (1) G. thunbergianum,
met var. thunbergianum en var. hirsutum in Oos-Rhodesi€, Transvaal, Swaziland, Natal,
Oranje-Vrystaat, Lesotho en Oostelike Kaapprovinsie; (2) G. capense, met ssp. capense In
die Suidwestelike Kaap en Oostelike Kaapprovinsie, ssp. namaquense (stat. nov.) in die
Westelike Kaapprovinsie, en ssp. garipense (stat. nov.) met var. garipense en ae
wittbergense (stat. nov.) in Transvaal, Natal, Oranje-Vrystaat, Lesotho en Ooste ne
Kaapprovinsie; (3) G. monticolum, ’n skaars spesies vanaf die Westelike Kaapprovinsie,
*Present address: Department of Botany and Microbiology, University of the Wit-
watersrand, Johannesburg.
Accepted for publication 1st November, 1977.
203
204 Journal of South African Botany
(4) G. bredasdorpense (sp. nov.) endemies aan die kalkbulte in die Bredasdorp-distrik; (5)
G. amatymbicum, ’n skaars spesies van die Oostelike Kaapprovinsie; (6) G. bussei, met
var. bussei en var. glabrum in Rhodesié; (7) G. scabrelloides (sp. nov.) in Suid-Transvaal,
Swaziland, Natal, Oranje-Vrystaat, Lesotho en Oostelike Kaapprovinsie; (8) G. scabrellum
in Oos-Rhodesié; (9) G. subvillosum met var. subvillosum en var. subglabrum (var. nov.),
(10) G. mucroniferum, met var. mucroniferum en var. dregeanum (stat. nov.) en (11) G.
undulatum (nom. nov.) in die Suidwestelike Kaap; (12) G. tomentosum in die Kaapprovin-
sie en suid Suidwes-Afrika (en Oranje-Vrystaat); (13) G. chloroionanthum in Oos-
Rhodesié; (14) G. spurium ssp. africanum, ’n skaars takson wat voorkom in die Transvaal,
Natal en Kaapprovinsie.
Nuwe chromosoomgetalle word gegee vir G. scabrelloides en G. capense ssp.
garipense var. wittbergense (n = 11, 2n = 22), en G. mucroniferum var. mucroniferum en
G. subvillosum var. subvillosum en var. subglabrum (n = 22, 2n = 44). Voorheen
gepubliseerde chromosoomgetalle in G. thunbergianum var. thunbergianum en var. hir-
sutum en G. capense ssp. garipense var. garipense (n= 11, 2n= 22) en G. tomentosum
(n= 22, 2n= 44) word bevestig deur aanvullende nuwe tellings.
INTRODUCTION
The Southern African members of the genus Galium are badly in need of
revision, the last comprehensive treatment of the genus being that of Sonder
(1865) who, in Flora Capensis, listed 18 taxa belonging to 14 species, 6 of which
were newly described by him. Since then only two short articles dealing with
South African taxa of the genus have been published (Phillips, 1917: description
of a new variety; Salter, 1937: discovery of dioecism in G. tomentosum). A few
more publications which primarily concern tropical East African species only
marginally touch problems encountered in South African taxa: Brenan (1953)
describes three new varieties of G. bussei and discusses their confusion with G.
stenophyllum; Ehrendorfer & Verdcourt (1973) discuss the relationships between
various tropical species, some of which extend into South Africa; Verdcourt
(1975) discusses the identity of G. spurium ssp. africanum (= ‘‘G. aparine’’) and
separates G. thunbergianum into two varieties.
The present publication not only covers South Africa (including Transkei),
South West Africa, Swaziland and Lesotho, but also Rhodesia,! where a number
of taxa centred in the tropics reach their southern limit of distribution. In order to
demonstrate the often obvious links between a number of tropical and Southern
African species this addition was considered necessary and useful.
The present paper offers a revision of the nomenclature and taxonomy of the
taxa in question, but also includes detailed information about morphology,
karyology, pollen measurements, as well as distribution, habitats and flowering
periods of the majority of taxa, although it needs to be stressed that this study
must by no means be considered complete, since information available for some of
the species still remains rather unsatisfactory.
" No collections seem to exist from Botswana and Mogambique (South of the Zambezi),
although particularly in the latter country some taxa are likely to occur in the mountain
ranges bordering Rhodesia.
The Genus Galium L. (Rubiaceae) in Southern Africa 205
MATERIAL AND METHODS
The present investigations were based on herbarium material borrowed from
the herbaria B, BOL, E, GOET, GRA, K, LISC, NBG, NH, NU, P, PRE, S,
SAM, SRGH, UPS, W, WIND, and WU (abbreviations after Index Herbariorum,
Holmgren & Keuken, 1974), and my personal collections made while in Southern
Africa between March 1976 and February 1977. Some of the material was
cultivated in a garden in Pietermaritzburg/Natal.
Karyological Investigations and Pollen Measurements
Material and methods as described in Puff (1976a).
Sem Investigations
Living or herbarium material was viewed with a Hitachi SSM 2 at an
accelerating voltage of 20 kV. Living material usually needed no preparation at all
(Figs 6c, 8e, 8d, for example), although sometimes results were only satisfactory
if it was first critical point dried and then coated with gold-palladium (Figs 8a, 8b,
for example). Herbarium material generally only required coating with gold-
palladium. Flowers taken from herbarium sheets were first boiled to restore their
original shape, then subjected to critical point drying, and finally gold-palladium
coated (Fig. 8c); this method cannot always be used because of unnatural
deformations as a result of this treatment.
Distribution Data and Maps
The ‘‘Degree Reference System’’ as outlined by Edwards & Leistner (1971) is
adopted. For greater accuracy, quarter-degree references are given whenever
possible. In the distribution maps one dot stands for all collections made ina
single quarter-degree square, whereby the actual position of any one collection
within the square is neglected. Degree references are also used for collections
from Rhodesia.
To establish Degree Reference Numbers the index of Southern African Place
Names (Leistner & Morris, 1976) was consulted. In the listings of collections the
spelling of place names has also been adjusted to that given in Leistner & Morris.
Place names used by earlier workers which, in the meantime, have changed, are
usually given in brackets after their modern equivalent (synonymy after Zoo-
Historical Gazetteer, Skead, 1973). To trace down exact localities of Ecklon &
Zeyher collections, Standérter-Verzeichnis (including Nachtrag) and Verglei-
chungen (Drége, 1847a, 1847b, 1848) were consulted. .
Distribution maps of taxa centred in tropical (East) Africa (Figs 12, 16, 18,
22): The extra-Southern African range of distribution of these taxa was compiled
from data given in Enumeratio Plantarum Aethiopiae Spermatophyta (Cufodontis,
206 Journal of South African Botany
1965) and Flora of Tropical East Africa: Rubiaceae I (Verdcourt, 1976), and from
herbarium material I have seen myself.
Altitudes given in feet were always converted to metres.
Descriptions
It should be noted that descriptions of taxa also occurring in tropical Africa
were drawn up from Southern African material only.
Stem diameters were measured in the mid-stem region unless stated otherwise.
In the measurements of leaf widths the reflexed margins were not taken into
account.
KARYOLOGY
Chromosome counts were determined from preparations of root tips, somatic
mitoses in young buds (walls of ovaries, filaments), pollen mitoses, and pollen
mother cell (PMC) meioses. Since the somatic chromosomes of Galium are rather
small (and therefore often difficult to count), an attempt was made to determine
chromosome numbers from clearly arranged metaphase I and II chromosomes of
PMCs whenever possible.
Fic. 1.
Somatic chromosomes of (a) G. thunbergianum (x = 11; 2n = 22); pollen mitosis II of (c)
G. capense ssp. garipense (x = n = 11); pollen mother cells of various diploid and
tetraploid Galium taxa (b, d—f): meta-/anaphase I of (b) G. thunbergianum (x = n = 11),
metaphase I of (d) G. scabrelloides (x = n = 11), metaphase II of (e) G. spurium ssp.
africanum (x = 10; n = 20) and (f) G. tomentosum (x = 11; n = 22). The bar represents
10 ym (b—f; for a the bar is 5 zm). Further explanations in the text.
The Genus Galium L. (Rubiaceae) in Southern Africa 207
All taxa investigated (with the exception of G. spurium ssp. africanum) have a
chromosome base number of x = 11, the most common base number in the genus.
The tetraploid G. spurium ssp. africanum is characterized by the derived base
number of x = 10. Within the x = 11 group diploids (G. thunbergianum, G.
capense ssp. garipense, G. scabrelloides) and tetraploids (G. subvillosum, G.
mucroniferum, G. tomentosum) were detected. Chromosome determinations for G.
thunbergianum var. thunbergianum and var. hirsutum, G. capense ssp. garipense
var. garipense, and G. tomentosum (Puff, 1977) are confirmed by additional
counts, the chromosome counts for G. subvillosum var. subvillosum and var.
subglabrum, G. mucroniferum var. mucroniferum, G. capense ssp. garipense var.
wittbergense, and G. scabrelloides are new:
Voucher Specimens for Chromosome Counts
number of
individuals
G. thunbergianum var. thunbergianum n 2n investigated
lng DOMZZIR2I (WNAU) 5 obo aoocens a0b6500b0 000 11
Ith (CURIE (ANAO)) ca coucsccagoso0G0D [00d jl 22 2
Jari OMMAS=3/b) (WWAW)) cesacnoccaneovsouco00ar 11 3
Additional counts are published in Puff (1977)
G. thunbergianum var. hirsutum
Pup 7OlO2—2 [1 (NUS WU) a. 11 22 6
Jt FOIA Hl (WKY) oo aceccconbo0 9. cdoo00c 11 4
JETT CUBIC LIU (\NAB)) aoc bocoaauoeobesdoonacGe DD 2
Puff 760418—I /4 (NU, WU) .........---+++---: 11 3
Puff 7612195 /1 (WU) (Fig. la, b) ......-...--- 11 22 4
Additional counts are published in Puff (1977)
G. capense ssp. garipense var. garipense
Puff 7612202 /3 (NU, WU) .........------+>>: 11 3
Ruff 76122031 (WU) :.--.------- + ee 11 22 3
PU OlZZ6—1 TaN) petaei ee ite ie 11 1
Additional counts are published in Puff (1977)
G. capense ssp. garipense var. wittbergense
Puff (OIZI8=1/Sby(NU, WU) sean 11 22 5
Puff 761219-1 /2 (NU, WU) .....--------+++-:> 11 3
Puff 7612192 /Ic,d (WU) ......--+-200-0220 ee: 11 22 4
Puff 761219-6/1 (WU) ........--------2030- 11 2
Puff 761220-4 /1 (WU) .....-.--------2207022 11 3
Puff 761225-2 /2 (WU) (Fig. Ic) .....-----++-->> 11 22 5
Puff 761227-11 (NU, WU) ...---.-..---*+---- 11 4
G. scabrelloides
Pig 7OORI2 AU 2s (MW) yt says 0 oe 22 2
I) Js (\\ 1D) OOo SRE SETI ET c.22 I
Si) (INU, WAU) conemoonocces Hodes 11 22 5
Ail (Ux tOe WAG) cco ecmone coc dpe oco 11 22 4
LOTR Gi) MD Ns ONE Cte eras 22 1
1 Localities of voucher specimens collected by the author can be found in the ‘‘Collec-
tions’’ section of the respective taxa.
208 Journal of South African Botany
number of
individuals
n 2n investigated
Puff 760509-1 /1 to 1/5 (all WU).............. 22 8
2 VOOS!IO—2/S (NO) cococccccacacganccnooce 2) 2
Puff AOLI21—3)/3i (Wi) arrest eeoreee 11 22 3
LP; WOIUAS-BO) (WNW) occccog00050 500 9000c006 11 2
i YOLAWLT/| (ONIW, WAD) coccocvavecacgvvcas 11 22 2
JONG VONAHOAYH (NW YWAD) ob00000000000cd000¢ 11 22 3
LG UOLAZLOAS (WD) aoccccocsceagvcadvve0Ke 22 1
24. ISN Wi) ate nch eee ere eee 11 22 5
Vif VOHZZIS ll (NW, WAG) cocccccoocoocs0cees 11 2)
25) TONE Wi) ie ae sacte ee cre eae 11 3
=A) (NW: Wi): abe eee eee iil 2
Lif LHIZ2ZS ANP (NNW WAG) ooccccoccovccccvces 11 1
LAN DONZZOANID (ONO) coccccsccccscscccc0ccc: 11 22 D;
IPF PIOUS (WNW) C8 UC) coccccoovcc0cc. 11 3
G. subvillosum var. subvillosum
JAI USOLOSO/2 (NNW, WAG) coccccccaccncnccuce 22 4
Pi AOOQUSD fll (NNW; WAWD) coccoccccccccccccee 22 44 5
G. subvillosum var. subglabrum
Ln MCOLOSSDPiLll WNW, WAD)) coccccesccccsenccs Wp) 44 3
LAV USOL2O=SYYll (NNO, WAG) wc ocececasscavcacce 22 4
SV; ASOLLOD EF (NI, WAU) occ cscecccccacdouace 2D 44 3
G. mucroniferum var. mucroniferum
LNT OOLOSHOI) (WO) ooccocccaccccoeoscascen 2D; 44 3
G. tomentosum
len LEOLMOA il (NW, WW) 2 ooceccaccocccoece c.44 2
Puff 7609131 /I (NU, WU) 6 (Fig. 1f).......... 22 3
Slip) (NW, WU) G5 2 foccsccccccc0ce 22 44 5
AY ONIW, WAU) S oaccoacoo00bcbe 22 D
LN LOOM il ONMW, WAY) @ cooccccscccarccce 22 4
Leh SOHAIL (UNM, WAU) @ 5 2 aosccccccsoccce LD 44 4
Fagerlind (1937), material of unknown origin (as
“G. Gyna@7ine Wowmly,”’)) ococcococvcccabeaoce 44
Additional counts are published in Puff (1977)
GASDUTIATESS DHIG/GI GAL a ne 20 40
Counts of material from the Southern Transvaal
and Natal are published in Puff (1977). Fig. le is
from Puff 770102-3 /1 (NU, WU).
POLLEN MEASUREMENTS
Pollen investigations were carried out on 305 individuals belonging to 12 taxa;
chromosome numbers of 113 of these were known. The pollen quality, although
more or less variable, in general is fairly good (most of the grains well developed
and three-nucleate; few grains dead, ‘‘giants’’ or ‘‘dwarfs’’). Only in G. scabrel-
loides, G. tomentosum and G. spurium ssp. africanum were abnormal grains found
The Genus Galium L. (Rubiaceae) in Southern Africa 209
more frequently. The occurrence of abnormal grains in these taxa, however, seems
to vary even within the population level (particularly in G. tomentosum).
It is obvious from Figs 2 and 3 that the average pollen diameters of diploids
are smaller than these of tetraploids. Within the diploids, the pollen of G.
thunbergianum is, on the average, smaller than that of the other taxa. Taking into
account the fact that within a given ploidy level pollen diameters are + constant in
closely related taxa (cf. Puff, 1976a and literature cited there), it can be concluded
that G. capense ssp. capense and ssp. namaquense (no chromosome numbers
known) are diploid, like G. capense ssp. garipense.
In tetraploids, pollen sizes of a particular species seem to have a much larger
range than those of diploid species (for example, G. spurium ssp. africanum: 22,4
pm to 26,5 wm—a range of 4,1 wm). G. subvillosum, the closely related G.
mucroniferum, and the presumably polyploid (tetraploid?) G. undulatum have
much larger grains than the other tetraploids.
The pollen grains of the tetraploid G. spurium ssp. africanum (Fig. 3c) are, on
the average, larger than those of the North hemispheric G. spurium f. spurium and
f. vaillantii (DC.) R. J. Moore (diploid, n = 10; 18-24 wm: Moore, 1975): this
once again seems to prove that in related taxa pollen becomes larger with
increasing ploidy level. The average pollen diameters of G. spurium ssp. afri-
canum, however, are smaller than those of the hexaploid and (at least morphologi-
cally) closely allied G. aparine f. aparine and f. intermedium (Bonnet) R. J.
Moore (n = 33; 25-31 um: Moore, 1975).
Pollen measurements of plants belonging to the “‘G. aparine-spurium com-
plex’’ would indicate the occurrence of (at least some) diploids: for a discussion of
this problem refer to G. spurium ssp. africanum *‘Critical Remarks’”’.
: ii rT q{2L_ID)
| pe eon ) L
aS. aes = : :
ee ee
Fic. 2.
Average pollen diameters in wm of G. crea ‘
ssp. namaquense (c), and ssp. garipense (d); G. scabre {
ablillosion (g); G. mucroniferum (h). Individuals investigated karyologically: black.
Vertical scale is numbers of individuals.
a); G. capense ssp. capense (b),
loides (e); G. undulatum (f); G.
210 Journal of South African Botany
Fic. 3.
Average pollen diameters in wm of G. chloroionanthum (a); G. tomentosum (b); G.
spurium ssp. africanum (c); ‘‘G. spurium-aparine complex’’ (d). Individuals investigated
karyologically: black. Vertical scale is numbers of individuals.
It should also be noted that, in addition to the correlation between pollen
diameter and ploidy level, there also seems to be a size correlation between pollen
mother cells and chromosomes: As can be seen in Fig. 1, pollen mother cells of
tetraploids tend to be somewhat larger than those of diploids (compare Fig. le, f
and b, d), while, on the contrary, chromosome size somewhat decreases with
increasing ploidy level (particularly obvious in Figs 1d and f). This phenomenon
has already been observed in closely allied species complexes (Puff, 1976b and
literature cited there) but, as can be seen here, it also seems to hold true for
unrelated or at least not closely related taxa of a genus.
MorPHOLOGY
Habit, stem and branching
With the exception of G. spurium ssp. africanum (annual) all Southern African
taxa are perennials with an often more or less woody base or rootstock/rhizome. In
G. tomentosum the lower half of the stem is often very woody (up to 20 mm in
diam. near the base), but this is never so pronounced in the other species. The
length of the stems ranges from 2,5(3) m (G. tomentosum) to ca. 100 mm (G.
thunbergianum, for example), their diameters (in the mid-stem region) from ca.
3(4) mm (G. tomentosum) to less than 1 mm (G. amatymbicum). Stems are
sometimes more or less terete near the base (G. capense, for example), but always
4-angled (collenchyma strands) upwards. Stems may be erect or ascending (G.
capense ssp. garipense, for example), caespitose (G. amatymbicum) or scrambling
or straggling over surrounding vegetation (G. tomentosum, for example). They are
usually green, but sometimes purplish or purple (anthocyanin. G. subvillosum, for
example). Internode lengths vary from 80-100 mm (G. chloroionanthum, G.
tomentosum, for example) to ca. 1—5(6) mm (G. bredasdorpense).
The branching is often very variable (sometimes even within a single species)
and is probably—as demonstrated for the non-South African Galium sect. Apa-
rinoides (Puff, 1975)—influenced by environmental factors. Depending on the
angle between main axis and lateral branches and the length of the latter, the shape
of the plants is broadly pyramidal (long, more or less horizontal lateral branches
The Genus Galium L. (Rubiaceae) in Southern Africa 2
which decrease in length towards the apex; G. scabrelloides, for example)
narrowly pyramidal (acute angle between axis and short lateral branches; G.
monticolum, for example) or (narrowly) cylindrical (short, more or less horizontal
lateral branches of approximately equal length; G. mucroniferum, for example).
AI
Fic. 4.
Stem portions of G. tomentosum (a), G. capense ssp. capense (b), and G. spurium ssp.
africanum (c) showing characteristic hair covering, a, c: X 20; b: x 30.
The indumentum of the stems can consist of densely set long, more or less
spreading whitish hairs (villous; G. tomentosum: Fig. 4a), short, spreading white
hairs (G. subvillosum, for example), short, curled white hairs (G. capense: Fig.
4b), more or less massive reversed prickles on the collenchyma strands (G.
spurium ssp. africanum: Fig. 4c, for example), or minute prickles (G. capense ssp.
garipense var. garipense, for example). In some taxa the stems are glabrous.
Leaves
Leaves and leaf-like stipules are arranged in whorls of 4 (G. thunbergianum),
6 (G. undulatum, for example), 6—8 (G. capense ssp. namaquense, for example),
or 6-10 (G. bussei, for example). They vary greatly in shape and length: middle
cauline leaves can be roundish-ovate, obovate, oblanceolate, ovate-lanceolate,
linear-lanceolate or linear, and are ca. 2 mm (CG. bredasdorpense) up to 35(40)
mm (G. chloroionanthum) long. Leaf shape and size can be quite variable within a
species (cf. G. tomentosum and G. thunbergianum: Fig. 5a, b), and even a
population (cf. Fig. 5a) most probably due to environmental influences [as in the
Central European G. mollugo group (Zimmermann, 1932) and Galium sect.
Aparinoides (Puff, 1975)]. In the majority of taxa, however, at least the leaf
DD Journal of South African Botany
1099400 100000
IGS:
Variation in size and shape of middle cauline leaves of G. tomentosum (a; *‘x’’ denotes
leaves taken from one population), and G. thunbergianum (b). The bar represents 10 mm.
shape remains more or less constant. The leaves of all taxa are 1-nerved with the |
exception of those of G. thunbergianum (3-nerved). In all South African taxa the {
apex of the leaf has a more or less distinct hyaline point or mucro (cf. G.
scabrelloides, Fig. 6c). The leaf margins are often strongly revolute (cf. G.
capense ssp. namaquense: Fig. 6b). Leaf texture ranges from more or less
membranaceous (G. amatymbicum) to more or less coriaceous (G. tomentosum,
for example).
Fic. 6. |
Leaf portions of G. subvillosum var. subvillosum (a), G. capense ssp. namaquense (b), and
G. scabrelloides (c, d) showing characteristic hair covering. a—c: X 20; d: x 120.
The Genus Galium L. (Rubiaceae) in Southern Africa M3}
The following types of indumentum occur: (coarse) reversed prickles on the
margin (G. scabrelloides: Fig. 6c, G. tomentosum: Fig. 6d) and/or white, +
straight spreading hairs on the lamina (G. subvillosum: Fig. 6a, G. scabrelloides:
Fig. 6c, for example), or small, + forwardly directed prickles on margin and
lamina (G. capense ssp. namaquense: Fig. 6b). In some taxa (G. thunbergianum
var. thunbergianum, for example) the leaves are always glabrous.
Synflorescences
In Galium the whole inflorescence system is a monotelic synflorescence sensu
Troll (1964): main axes and all other branches (axes) terminate in a flower. To
avoid too complex descriptions of the synflorescences all lateral ‘‘inflorescences’’
arising from the main axis are here called paracladia (cf. Fig. 7: pc).
Fig. 7 represents the full spectrum of synflorescence types found in Southern
African Galium species: Fig. 7a depicts the most common type (in G. capense, G.
scabrelloides, G. tomentosum and others), in which the paracladia are many-
sf
lox
Cc d e f
Fic. 7.
Schematic illustration of synflorescence types. Only three nodes of the ue ae
region are shown; it is taken into account that not always two, oneae ie wae oe
paracladium (pc), depending on the species, is produced at one node. br: zee) a
of bracts; ma: main axis; sf: synflorescence; wh: whorl of foliage leaves. Further exp
tions in the text.
214 Journal of South African Botany
flowered and the flowers are located on axes of up to the fourth order (in
taxonomic literature frequently somewhat incorrectly termed ‘‘axillary panicles’’).
The type illustrated in Fig. 7b (in G. thunbergianum, G. undulatum, for example)
has somewhat fewer-flowered paracladia with flowers only on axes of the second
and third order. Types transitional between those of Fig. 7a and b may occur
(owing to environmentally controlled reduction in flower number?), and are fairly
common particularly in G. capense. The types illustrated in Fig. 7c and d have
paracladia further reduced: they consist of only one of two axillary or terminal
groups of three or two flowers (G. subvillosum and G. mucroniferum, respect-
ively). In the type illustrated in Fig. 7e the paracladia are very short and have a
whorl of leaf-like bracts and three to one flowers only (G. bredasdorpense).
Finally, Fig. 7f shows a type in which the paracladia are reduced to the terminal
flower: in G. spurium ssp. africanum and G. chloroionanthum the one-flowered
paracladia are always short (in the latter there is always a bract just below the
flower), while in G. amatymbicum the single flowers may also be located on more
or less long paracladia.
It should be noted that the above reduction series in the synflorescence region
does not reflect direct phylogenetic relationships of the taxa mentioned as
examples. Such reduction series have evolved independently from each other in
several other Galium species.
The shape of the synflorescence may vary considerably depending on the
length of the paracladia and the angle between paracladia and the main axis: cf.
“Habit, Stem and Branching’’.
Pedicel length can range from (0,5)1(2) mm (G. amatymbicum, for example)
to 10(20) mm (G. mucroniferum, for example); in 2 plants of G. tomentosum they are
much elongated and up to 60(90) mm long. They may be more or less thick and
firm (G. spurium ssp. africanum, for example) or thin, more or less filiform (G.
amatymbicum), straight and more or less divaricate (most taxa) or more or less
arcuate (G. spurium ssp. africanum, for example) in fruit. In the majority of taxa
they are glabrous; in G. spurium ssp. africanum they are often beset with reversed
prickles, in G. subvillosum var. subvillosum, for example, with more or less long,
whitish hairs; in G. tomentosum they are villous.
The peduncles are usually about as long as the pedicels and correspondingly
hairy.
Flowers
All Southern African taxa with the exception of the dioecious G. tomentosum
(Fig. 8a, b: 2, note rudimentary stamina; Fig. 8c: ¢) have hermaphrodite flowers.
The 4-merous corolla is always rotate, glabrous (most taxa) or somewhat hairy
outside (G. scabrelloides, cf. Fig. 8d, or G. thunbergianum var. hirsutum, for
example). The corolla lobes are more or less ovate to triangular, about as long as
wide (G. chloroionanthum) or (much) longer than wide (all other taxa), acute or
The Genus Galium L. (Rubiaceae) in Southern Africa 215
acuminate, but never obtuse. Corolla colour ranges from brilliant yellow, yell
pale and creamy yellow, yellowish-green to greenish-white or tea (s ‘ yee
colour is variable even within a species: G. thunbergianum, for oan ae
none of the Southern African taxa have pure white corollas. In G. ee is
FIG
Flowers of G. tomentosum (a, b: 2; c: 3), G.
d, e: X 20; b, f: x 60; c: x 80. Explanations in the text.
5 te :
cabrelloides (d), and G. subvillosum (e, f). a,
216 Journal of South African Botany
corollas may occasionally have a purple tinge (sun-exposed specimens?; corollas
normally creamy yellow). The calyx is always completely reduced. The two-lobed
disk (superimposed on the 2-carpellate inferior ovary) is usually more or less
massive (cf. Fig. 8f). The 4 stamina are nearly as long as the corolla lobes (G.
mucroniferum, G. subvillosum: Fig. 8e) or much shorter (less than half as long as
the corolla lobes: G. scabrelloides: Fig. 8d; even shorter in G. tomentosum G: Fig.
8c); the anthers are usually more or less elliptic. The styles are fused to the middle
(or at least 1/3), the free ends often arch downward; the stigmas are always
capitate.
Fruits
The fruits consist of two mericarps which are globose to subglobose (most
Southern African taxa, cf. Figs 9 and 10 a—d) or more or less reniform (G.
tomentosum: Fig. 10f). They are always dry (never fleshy). Their surface can be
glabrous (usually more or less granulate: G. tomentosum, Fig. 10e, for example) or
covered with white, stiff, hooked hairs which are tuberculate (on a distinct
elevation of the skin tissue: cf. Moore, 1975; G. thunbergianum, Fig. 9a, b, for
example) or non-tuberculate (G. spurium ssp. africanum, Fig. 9c, d, for example;
wrinkles in the skin, however, may occasionally simulate tubercles). In G.
chloroionanthum (Fig. 9e, f) the hooked hairs are always brown and appear to be
flattened at least on herbarium material; no fresh material was available. The
mericarp surface can also be covered with (more or less long) straight, white
spreading hairs (G. scabrelloides, Fig. 10a, b: tuberculate), very short, straight
hairs (G. subvillosum, Fig. 10c, d), or short, more or less curled hairs (G. capense
ssp. garipense; occasionally in G. tomentosum, cf. Fig. 10f). Sometimes only one
carpel develops into a fruit. In this case the hairs are much more closely spaced on
the aborted carpel than on the fertile one (cf. Figs 10c and 10d): obviously the
hairs are initiated at a very early stage of development and become more widely
spaced as a result of fruit diameter increase during development.
G. tomentosum has mericarps with three distinct grooves (cf. Fig. 10e), a
feature absent from all other Southern African species.
Mericarp size (diameter of a single mericarp) ranges from (0,7)0,8—1(1,2) mm
(G. amatymbicum) to (2)2,5—3,5 mm (G. undulatum, for example).
POLLINATION BIOLOGY
All Southern African taxa with the possible exception of the annual G. spurium
ssp. africanum (autogamous?) appear to be entomogamous. Field observations are
fragmentary, but small insects (small flies, not precisely identified) were at least
observed on flowers of G. tomentosum (6 and 2 plants), G. scabrelloides and G.
capense ssp. garipense. The flowers of the last two were found to be strongly
scented. Flowers of G. thunbergianum, G. capense ssp. garipense, G. scabrel-
loides, G. subvillosum and G. mucroniferum that were investigated in the field
The Genus Galium L. (Rubiaceae) in Southern Africa : DAY
Fic. 9.
Fruits of G. thunbergianum (a, b), G. spurium ssp.
thum (e, f). a, c: X 25; d, b: X 100; e: x 30;
africanum (Cc, 4d), and G. chloroionan-
f: x 120. Explanations in the text.
218 Journal of South African Botany
Fic. 10.
Fruits of G. scabrelloides (a, b), G. subvillosum var. subvillosum (c, da), and G.
tomentosum (e, f). a: X 30; b: X 200; c: x 10; d: x 30; e, f: x 20. Explanations in the
text.
The Genus Galium L. (Rubiaceae) in Southern Africa 219
ee
|
|
2
|
Fic. 11.
Schematic representation of mechanism preventing self-pollination in G. subvillosum.
Further explanations in the text.
were found to produce nectar in profusion; only in G. spurium ssp. africanum do
the tiny disks apparently produce very little nectar. G. capense ssp. garipense, G.
scabrelloides and G. thunbergianum have slightly protandrous flowers. In G.
mucroniferum and G. subvillosum, however, protandry is particularly pronounced
(Fig. 11): pollen is released as soon as the flower opens (Fig. lla). After one or
two days the filaments begin to move into a more or less horizontal position; at
this stage stigmas are not yet fully developed, and the styles have not yet
elongated (“male state’’ of the flower, Fig. 11b; also see Fig. 8e, f). After
approximately three days the styles slowly begin elongating (Fig. 11c); four to six
days after the opening of the flower the styles have reached their full length, and
the stigmas are ready for pollination (“‘female state’’ of the flower). Meanwhile
the filaments have bent down below the level of the corolla lobes (the anthers by
now are empty or almost empty; Fig. 11d).?
Similar mechanisms promoting allogamy have already been recorded for other
Galium species by Miiller (1873, Fig. 135: G. verum) and Kirchner (1911, Fig.
37: G. mollugo).
DELIMITATION OF THE GENUS
The genus Galium L. belongs to the tribe Rubieae. The only other genus of
this tribe represented in Southern Africa is Rubia with the taxa R. cordifolia L.
ssp. conotricha (Gandoger) Verdc., R. petiolaris DC., R. horrida (Thunb.) Puff
(recently transferred from Galium: Puff, 1978), and the introduced R. peregrina L.
(according to Sonder, 1865; no specimens seen by me). It is distinguished from
Galium (as far as Southern African material is concerned) by much longer leaves
with well developed petioles (excluding R. peregrina), S-merous corollas and
more or less fleshy fruits.
TAXONOMY
Galium L.”
Perennial herbs or annuals. Stems more or less distinctly 4-angled (sometimes
more or less terete near the base), glabrous or beset with hairs or reversed prickles.
Leaves and leaf-like stipules in whorls of 4-10, epetiolate. Synflorescences many-
1 Based on observations of plants cultivated in a garden in Pietermaritzburg. Results were
consistent in all plants examined. ; oe ‘
2 Description valid only for the Southern African members of the genus.
220 Journal of South African Botany
to few-flowered. Flowers perfect, or imperfect (plants dioecious); corollas 4-
merous, rotate, yellow to whitish; calyx absent; stamina 4, as long as or shorter
than the corolla lobes; ovary inferior, stigmas capitate; fruits consisting of two
globose, subglobose or more or less reniform mericarps, dry, glabrous or with
hooked or straight hairs.
KEY TO THE SPECIES AND SUBSPECIES
ow Nn =
=
=
oO
©
Middle cauline leaves in whorls of 4, 3-nerved.............. 1. G. thunbergianum Nor Sy i;
Middle cauline leaves in whorls of 6—10, Il-nerved............................ 2 Desi
Synflorescences very reduced, paracladia with one or two groups of 3—1 flowers .. .3
Synflorescences + extensive, paracladia many-flowered ........................ 8
. Leaves large, up to 38(45) mm long and (12)15 mm wide, with reversed prickles
ONithesmManeansspiUitspwathwOOKe Cals erie eee eens nee ete Pe
Leaves small, up to 14 mm long and 3 mm wide, without prickles on the margins; =a proce) )
. Flowers imperfect, plants dioecious, peduncles and pedicels villous ...........
. Leaves with densely set reversed prickles on the margin; pedicels hairy with +
ELE VEISed MLICkles spedicelsselabrouslon ScablOUS se eee eaten nena 2
. Longest internodes (25)30—65(75) mm; mericarps typically hairy with straight,
. Leaves (18)20—30 mm long; from tropical East Africa extending into Rhodesia .
TOUS MEY OUS Or Valin aeIeN, MOE NCO Gl EVES 2. oe noo pon occ oo scceseoensseooe
Fruit hairs brownish, ca. 1 mm long, flattened, very densely spaced; pedicels 0,5—
Memimulone.< sevens Set cel One Ee 13. G. chloroionanthum <p
Fruit hairs white, less than 0,5 mm long, not flattened, well spaced; pedicels
(GLASS) Mv WONG osaccoccodvacoodsasccucove 14. G. spurium ssp. africanum TH We
Leaves large, up to 14 mm long and 3 mm wide; longest internodes 10—45(55) io 7)
mm; fruiting pedicels up to 10(13) mm long; corolla (2,5)3—4 mm in diameter ..... 6
Leaves small, up to 5(6) mm long and 1,2 mm wide; longest internodes 1—10(15) - *~**2%2,/4
mm; fruiting pedicels up to 1(2) mm long; corolla 1,5—2 mm in diameter ......... as
Paracladia with 3—6 flowers arranged in groups of 3; fruiting pedicels (3)4—6(8)
MIMNONB Sh kieces Steen. Sua vis ae eh cers ieee ences Smee uc Ren eene 9. G. subvillosum 5” Ca,
Paracladia with 2—4 flowers arranged in groups of 2; fruiting pees (4)6—10(13)
TMIMULOM Be 2 sais es keys Sel ately Cos iee eek xk ne Oe 0. G. mucroniferum £4) GC,
Longest internodes 1—5 6} mm; leaves up to 2,5 mm long, aaa paracladia
3 =I flowered. Gran. he con eeepc emw reg crane ere eee 4. G. bredasdorpense }“4«j
Longest internodes 8—10(15) mm; leaves up to 5(6) mm, with+ long, white » i
spreading hairs; paracladia only 1-flowered.................. 5. G. amatymbicum / eA
SE RE RT iti oer 8 ENS SEE RARE hood Rug ocohehiove 12. G. tomentosum 4, C|
Flowers perfect, peduncles and pedicels glabrous, scabrous, or with a few, + Capty
Stralghtawhiteshaits sbUteneVeTavill OU Smee ie rer nen eine 9 i
Leaves obovate to obovate-oblong, (6)8—10(11) mm wide, often glaucous, always
in whorls of 6; flowers with very short stamina (anthers almost sessile) ........
a lbogi se sag te asada cb eS aay he 11. G. undulatum © >-"
Leaves linear, lanceolate or (narrowly) oblanceolate, (0,6)0,8—4(5) mm wide, Be A
never glaucous, in whorls of 6—8—10; flowers with stamina ca. half as long as the i
fof) (0) U1 (0) of le ie arr OR hee aoe ee cue coduoes 10 |
long, isiraight white|hairs| om elabrousi.. sera eee eee eee eee Il
Leaf margins glabrous, with + long, straight white hairs or only very few, small
white hairs; leaves up to 4(5) mm wide; South Africa, Lesotho, Swaziland ..... ot al
NOR ae eee yeni th carder ateechire:- cian ammicin eileen Be Sie 7. G. scabrelloides Nt Sty
Longest internodes 20-25 mm; mericarps glabrous; leaves not wider than 2(2,5)
mm; from tropical East Africa extending into Rhodesia........... 8. G. scabrellum /~»-
mfigi ave ae 9. } uavinmtnce: Woe) ele Vai scouslino, wee ane Dely Soe se ee ete 6. G. bussei Trop Ne
Reaves' (3)5—20(23) mmilong;SouthyAtiica,sleesotho) see ane aaa eee 13} |
The Genus Galium L. (Rubiaceae) in Southern Africa Mp2
13. Stems densely covered with short, spreading hairs; leaves 0,6—0,8(1) mm wide
surfaces densely covered with + long, white spreading hairs; corolla (1,5)1 Te
2233) Taito Th, GHENT Cie 5 ais ears G enn dete De epee eet ite co tear: 3. G. monticolum
Stems glabrous, with minute prickles or very short, curled whitish hairs; leaves
(0,3)0,5—2(2,5) mm wide, surfaces smooth or with small, forward-directed
prickles or short straight hairs; corolla 2—3,5(4) mm in diameter ................ 14
14. Leaves with small, forward-directed prickles or short, straight hairs on the
surfaces; pedicels scabrous (or subglabrous) ....°. .2B. G. capense ssp. namaquense
Leaves glabrous or with a few (réversed prickles on the margin; pedicels always - =
PLADLOUSWaP Ey s Pats yaycests chs, Sei va his ed owidaratae Shahi aap evoraer-wcka ines sowie: 15
15. Mericarps glabrous; leaves up to 12(15) mm long...... 2A. G. capense ssp. capense
Mericarps + densely covered with short, curled whitish hairs (rarely subglabrous);
leavesmipntor20@3)immiloneie esp -eae sae eae: 2C. G. capense ssp. garipense
1. G. thunbergianum Eckl. & Zeyh., Enum. Pl. Afr. Austr.: 369 (1836). Type:
South Africa, ‘‘Ceded Territory’’ (= SE. Cape Prov.), Katriviersberg, Ecklon
& Zeyher 2321 (S!, SAM!; NU, WU: photos!).
G. rotundifolium sensu Thunb., Fl. Cap. 1: 551 (1813), non L.
G. rotundifolium sensu Sonder in Fl. Cap. 3: 39 (1865), non L.
G. rotundifolium L. var. hirsutum Sond. in Fl. Cap. 3: 39 (1865). Type: South
Africa, Cape Prov., Masson in herb. Thunberg sub no. 3354 (UPS!,; NU, WU:
photos!).
G. rotundifolium L. var. normale O.Ktze., Rev. Gen. PI. 3,2: 120 (1898).
Type: South Africa, Natal, Van Reenen’s Pass, 1800 m, O. Kuntze s.n. (K!).
G. dayscarpum Schweinf., Beitr. Fl. Aethiop.: 135 (1867). Type: Ethiopia,
Simien, Debra-Eski, Schimper (B).
G. biafrae Hiern in Fl. Trop. Afr. 3: 245 (1877). Types: Fernando Po, Mann
605 & Cameroun Mtn., Mann 1284 (both K).
G. natalense Rouy in Fl. Fr. 8: 9 (1903), adnot. Type: South Africa, Natal,
Drakensberge, Van Reenen Pass, Medley Wood 5562 (LY; K: photo).
Perennial with extensive rhizome. Stems climbing, suberect or procumbent,
(80)100—400(600) mm long, with + few lateral branches; 0,5—1(1,5) mm in diam.,
distinctly 4-angled, glabrous or with short, + spreading white hairs. Longest
internodes (15)25—50(65) mm. Middle cauline leaves in whorls of 4, 3-nerved,
(7)10—18(22) x (3,5)4—-8(10) mm, (roundish-)ovate, elliptic or + rhombic, +
acute or with a short hyaline point at the apex, + cuneate at the base, glabrous or
with short, + spreading hairs on both surfaces and on the flat margins. Synflores-
cences broadly pyramidal to + cylindrical, paracladia + many-flowered, ultimate
branches with 1—2 minute bracts;’ pedicels 1-3(5) mm, slightly elongating after
anthesis, glabrous or hairy, + divaricate in fruit. Flowers slightly protandrous,
4-merous; corolla (1)1,5—2,5(3) mm in diam., rotate, often with a few short hairs
outside, greenish, greenish-white, pale yellow, or yellow, lobes longer than wide,
+ ovate; stamina ca. half as long as the lobes, filaments + filiform, anthers
elliptic; styles ca. 1/, to '/, fused. Fruits dry, densely covered with white
tuberculate hooked hairs; mericarps subglobose, each (0,8)1—-1,5 mm wide.
222 Journal of South African Botany
Two varieties recognized:
la. var. thunbergianum
G. natalense Rouy in FI. Fr. 8: 9 (1903), adnot. Type: South Africa, Natal,
Drakensberge, Van Reenen Pass, Medley Wood 5562 (LY; K: photo).
G. rotundifolium L. var. normale O.Ktze., Rev. Gen. Pl. 3,2: 120 (1898).
Type: South Africa, Natal, Van Reenen’s Pass, 1800 m, O. Kuntze s.n. (K!). |
Stems, leaves, peduncles and pedicels glabrous or nearly so (occasionally a
few hairs at some of the upper nodes or along the nerves of the leaves).
Chromosome Number: n = 11, 2n = 22.
Habitat: In forest margin vegetation (but rarely in forests), on stream banks,
under rock overhangs, or in rocky grassland. Generally in damp to wet, + rich soil
and sheltered, + shady locations. Ca. 900—2 350 m.
Flowering Period: December to March (April).
Distribution (maps, Figs 12, 13): In the mountains of West Kenya and
Ethiopia, and Southern Africa (East Rhodesia, East and South Transvaal, Swazi-
land, Natal, Lesotho, Orange Free State and Eastern Cape Province).
1b. var. hirsutum (Sond.) Verdc. in Kew Bull. 30: 326 (1975).
G. rotundifolium L. var. hirsutum Sond. in Fl. Cap. 3: 39 (1865). Type: South
Africa, Cape Prov., Masson in herb. Thunberg sub no. 3354 (UPS!; NU, WU:
photos!).
G. rotundifolium sensu Thunb., Fl. Cap. 1: 551 (1813), non L.
G. dayscarpum Schweinf., Beitr. Fl. Aethiop.: 135 (1867). Type: Ethiopia,
Simien, Debra-Eski, Schimper (B).
G. biafrae Hiern in Fl. Trop. Afr. 3: 245 (1877). Types: Fernando Po,
Mann 605 & Cameroun Mtn., Mann 1284 (both K).
Stems (mainly angles), leaves, peduncles and pues = ae densely | covered
with short, ce ate spreading white hairs. ai easy)
Chromosome NTbSe m = iil, an = 2
Habitat and Flowering Period: As in var. thunbergianum.
Distribution (maps, Figs 12, 13): From tropical Africa (Sudan, Ethiopia,
Fernando Po, Cameroun: Cameroun Mtn., Zaire, Uganda, Kenya and Tanzania) to
Southern Africa, where it occurs in East Rhodesia, North, North East and South
Transvaal, Swaziland, Natal, Lesotho, Orange Free State and Eastern Cape
Province.
Critical Remarks: While var. thunbergianum is very much rarer than var.
hirsutum in tropical Africa (cf. Verdcourt, 1976), both varieties are more or less ©
equally well represented in Southern Africa. Although they were often found
growing side by side in the field, intermediate (+ hairy) forms were never
detected. The subdivision of the species into two varieties, therefore, seems
justified.
| The Génus Galium L. (Rubiaceae ) in Southern Africa 223
,
|
| feeb
|
|
|
224 Journal of South African Botany
G. thunbergianum is very variable, particularly leaf size and shape (cf. Fig.
5b) and extent of the synflorescence may differ greatly. Since variation, however,
is considerable even within a single population and leaf and synflorescence
variation is not correlated at all, a further subdivision of the species does not seem
appropriate.
G. thunbergianum 31
oO var. t. f
a var. hirsutum }32
P i=] var. t. & var.h. |
33
A G. scabrellum |
| x
4 [cae ee ||
22 23 24 25 26 27 28 29 30 vu 32 33 34 35 36 37
Fic. 13.
Distribution of G. thunbergianum var. thunbergianum and var. hirsutum, and G. scabrel-
lum in Southern Africa.
The Genus Galium L. (Rubiaceae) in Southern Africa 225
COLLECTIONS
var. thunbergianum
RHODESIA— 1832: Inyanga distr. Stapleford, Nuza Slopes, ca. 1 830 m
(-DB), Wild 5706 (LISC, SRGH).
TRANSVAAL—2530 (Lydenburg): ca. 3 m from Whiskeyspruit on old Sabie-
Lydenburg rd., ca. 1 830 m (-BB), Meeuse 10331 (PRE, S); 5 m from Kaapse-
hoop to Ngodwana (-DB), Buitendag 823 (NBG).
——2729 (Volksrust): Farm “‘Highlands’’, Volksrust distr., ca. 1 700 m, Mogg
7521 (PRE).
——2730 (Vryheid): Wakkerstroom, ca. 1 830 m (-AC), Beeton 205 (SAM); Farm
““Oshoek”’ nr. Wakkerstroom, ca. 2 000 m, Devenish 799 (PRE); ‘‘South Hill’’
summit, ca. 2 300 m, Galpin 9830 (PRE); Kastrol Nek (-AD), Fitzsimmons & Van
Dam 26061 (PRE).
ORANGE FREE STATE—2827 (Senekal): Doorn Kop (-DD), Burke s.n. (K),
Zeyher 772 (K).
2828 (Bethlehem): Farm ‘‘Dunelin’’ nr. Fouriesburg (-CA), Potts 3088 (NBG,
PRE); Witsieshoek (-DB), Junod 17448 (PRE); Bester’s Vlei, ca. 1 500 m, Bolus
8171 (BOL), Flanagan 1866 (P, SAM).
—2926 (Bloemfontein): Thaba Nchu Mtn., Cave Kloof forest, ca. 2 100 m
(-BB), Roberts 2284 (PRE).
SWAZILAND—2631 (Mbabane): Hills N of Mbabane, ca. 1 380 m (-AC),
Compton 28691 (PRE, SRGH); Mbabane distr., Duiker Bush, ca. 1 500 m,
Compton 25596 (NBG).
NATAL—2731 (Louwsburg): Ngome (-CD), Strey 9557 (NH).
2828 (Bethlehem): Royal Natal National Park, Mont-aux-Sources, Pastures
Busch, ca.>1 830 m (-DD), Bayer & McLean 120 (PRE);—, Tugela valley and ~~“
Mont-aux-Sources, ca. 1 500-2 000 m, Humbert 14862 (P), 15015 (P).
2829 (Harrismith): E of Van Reenen, 1 700 m (-AD), Jacobsz 1562 (NBG);
Van Reenen pass, 1 800 m, Kuntze s.n. (K), Krook 2058 (S), Medley Wood 5662
(LY); Umlambonja valley (-CC), Marriott s.n. sub PRE 22654 (PRE).
—2929 (Underberg): Giant’s Castle Game Reserve, “‘Main Cave’’ (-AB), Puff
761221-2/1 (WU); Cathedral Peak Forest Reserve, catchment 9, ca 1 800—2 000
m, Puff 7603144 /2 (WU); Cathkin Park, floor of Ndema forest, ca. 1 370 m,
Galpin 11890 (PRE); Bushman’s R. nr. Dalton Bridge, ca. 1 400 m (-BB),
Wright, West & Acocks 3 (NH); Loteni Forest Reserve, S branch of Loteni R.
(BC), Wright 1475 (NU); Farm ‘‘Allandale’’, adjacent to Kamberg Nature
Reserve, ca. 2 100 m, Puff 761220-2/1 (WU); Bamboo Mtn. (-CB), McLean
681 (NH); nr. Bushmen’s Nek Police Post (-CC), Hilliard & Burtt 7985 (E, K,
MQ, NU, S); Bushmen’s Nek nr. Thamathu Cave, ca. 2 300 m, Hilliard & Burtt
se (E, K, NU, PRE); Garden Castle Nature Reserve, valley of Umzimkulu R.,
ca. 1 800 m, Hilliard & Burtt 7897 (E, K, MO, NU); summit of Nhlosane Mtn.,
226 Journal of South African Botany
nr. Dargle (-DB), Puff 761125-3/5 (WU); 3 m from Donnybrook on Ixopo rd., ca.
1 400 m (-DD), Edwards 3107 (NU, PRE); Polela, Medley Wood s.n. (SAM).
2930 (Pietermaritzburg): 6 m from Balgowan/Curry’s Post, ca. 1 350 m (-AC),
Moll 890 (NU, PRE); Keerom, Farm ‘‘Cottingham’’, ca. 1 380 m (-CC), Strey
8415 (NH, SRGH).
—3029 (Kokstad): Weza, Ingeli slopes (-DA), Strey 6305 (K, NH, NU);—,
Zuurberg, nr. Transkei Border Puff 760509 4/5 (WU).
LESOTHO—2828 (Bethlehem): Leribe, ca. 1 500—1 800 m (-CC), Dieterlen
146 (K, PRE, Ss, SAM); —, Lefi’s Kloof on NE slopes of Leribe Plateau, Phillips
972 (SAM); —, W slopes of Leribe Plateau, Phillips 749 (SAM).
——2927 (Maseru): Roma, ca. 1680 m (-BC), Ruch 1795 (PRE).
—2928 (Marakabei): Nsututse, ca. 2 440 m (-AA), Jacot Guillarmod 1242 (PRE).
Uncertain locality: ‘“S slope in ravine’, Schmitz 434 (PRE).
CAPE—3026 (Aliwal North): Aliwal North (-DA), Hepburn 279 (GRA).
——3028 (Matatiele): Potrivier Berg, ca. 1 680 m (-CD), Galpin 6650 (BOL,
GRA, K, NH, PRE, SAM).
—3029 (Kokstad): Invenyani nr. Cedarville (-AC), Bandert 200 (GRA).
——3225 (Somerset East): Bosberg, ca. 1 070 m (-DA), McOwan 1537 (BOL).
3226 (Fort Beaufort): ‘“Ceded Territory’, Katriviersberg (-D?), Ecklon &
Zeyher 2321 (S, SAM).
3326 (Grahamstown): Grahamstown (-BC), Glass 830 (NBG).
Uncertain locality: ““Cape of Good Hope’’, Cooper 2495 (K).
COLLECTIONS
var. hirsutum
RHODESIA— 1832: Inyanga distr., Inyangani Mtns. (-BB), Norlindh & Wei-
marck 5036 (SRGH); —, Inyanga Down (-BC), Norlindh & Weimarck 4740.
(SRGH); —, Chipungu Falls (-BD), Chase 4349 (SRGH). 7
—— 1832/1932: Umtali distr., Engwa, 1 980 m, Exell, Mendonca & Wild 306
(LISC, SRGH).
SRGH); Melsetter distr., near Bridal Veil Falls (-DD), Noel 2437 (LISC, PRE, SRGH).
TRANSVAAL—2230 (Messina): Hooge Veld, Pages Hotel (-AC), Rehmann
6862 (K).
—— 2329 (Pietersburg): Houtboschberg, 2 160 m (-DD), Schlechter 4387 (BOL,
PRE, S, SAM, W, WU), 47/8 (GRA, NBG, S, SAM, W, WU).
2330 (Tzaneen): Wolkberg Bosreserwe, path from Wolkberg to the
“Knuckles”’ (-CC), Puff 7701092 /3 (WU).
2430 (Pilgrim’s Rest): The Downs, ca. 1 200 m (-AA), Junod 4270 (PRE),
19973 (PRE); Pilgrim’s Rest, Black Hill, ca. 2 050 m (-DC), Galpin s.n. (BOL).
——2530 (Lydenburg): Mt. Anderson, 11,5 m W of Sabie (-BA), Strey 3042 (PRE,
SRGH); Belfast (-CA), Leedertz 9205 (PRE).
a
The Genus Galium L. (Rubiaceae) in Southern Africa DPT
——2627 (Potchefstroom): Krugersdorp (-BB), Jenkins 9263 (PRE).
2730 (Vryheid): Farm ‘*Oshoek’’ nr. Wakkerstroom, nr. Natal border (-AC),
Puff 770102-2/1 (NU, WU).
Uncertain or inexact localities: Warmbad distr., no locality given, Leipodt s.n.
sub PRE 42009 (PRE); Waterberg distr, W end of Waterberg Mtns., Farm
Groothoek, ca. | 800 m, Codd 3964 (PRE).
ORANGE FREE STATE—2828 (Bethlehem): Bethlehem (-AB), Potgieter
21850 (PRE), Bethlehem distr., Farm ‘‘Rondehoek’’, ca. 1 700 m, Scheepers
1763 (PRE).
Inexact locality: ‘“OFS’’, Cooper 2497 (E, K).
SWAZILAND—2631 (Mbabane): Hills N of Mbabane, ca. 1 380 m, Compton
28691 (NBG).
NATAL—2729 (Volksrust): nr. Charlestown, ca. 1 500-1 800 m (-BD), Mea-
ley Wood 5566 (GOET); Farm “*Glen Atholl’’ nr. Charlestown and Volksrust, ca.
1 800 m Smith 5693 (PRE); Majuba, Rogers 3310 (PRE).
——2730 (Vryheid): Vryheid distr., Farm Kambula, Gerstner 4632 (PRE).
——2828 (Bethlehem): Royal Natal National Park (-DD), Mont-aux Sources, Steyn
1061 (NBG), ca>1 800 m, Bayer & McClean 120 (PRE), ca. 2 000 m, Mogg 4247
(PRE), ca. 2 150 m, Sim s.n. sub PRE 42007 (PRE); —, Tugela valley and
Mont-aux-Sources, ca. 1 600 m, Humbert 14846 (P).
—2829 (Harrismith): Olivershoek pass, ca. 1 500 m (-CA), Wood 3558 (BOL, E,
NH, SAM), Strey 9526 (NH); Cathedral Peak area (-CC), Esterhuysen 10232
(BOL).
—2831 (Nkandla): Ulundi, ca. 1 500-1 800 m (-AD), Evans 486 (NH).
——2929 (Underberg): Giant’s Castle Game Reserve, nr. ‘‘Barne’s Shelter’
(-AB), Puff 761212-1/2 (WU); Cathedral Peak Forest Reserve, Forest Research
Station, catchment 3, ca. 1 900 m, Killick 1358 (BOL, NH), —, catchment 9, ca.
1 800-2 000 m, Puff 76031-/ /1,—2/1 (WU); Cathkin Peak, ca. | 400 m, Galpin
s.n. (BOL); nr. Champagne Castle Hotel, ca. 1 650 m, Acocks 10067 (NU, PRE);
Bushman’s R, nr. Dalton Bridge, ca. 1 400 m (-BB), Wright, West & Acocks 3 ~
(BOL); South Downs, ca. 1 500-1 800 m (-BB), Evans 514A (NH); Game Pass
Farm, Gladstone’s Nose ridge, at ‘‘Game Pass’’ bushmen shelter, ca. 1 800 m
(-BC), Puff 760418— /4 (NU, WU), —//5 (NU, WU); Farm Allandale, adjacent to
Kamberg Nature Reserve, ca. 1 900-2 000 m, Puff 761220-1/1 (WU); Kamberg,
ca. 2 000-2 100 m (-BD), Wright 1784 (NU), 2/16 (NU), Puff 761219-5/1 (WU);
Bamboo Mtn. (-CB), Grice s.n. (NU); Bushmen’s Nek, nr. Thamathu Cave, ca.
2 300 m (-CO), Hilliard & Burtt 8922 (E, K, NU, PRE); Garden Castle Nature
Reserve, valley of Umzimkulu R., ca. 1 800 m, Hilliard & Burtt 7897 (E, K, MO,
NU); Castle Gardens, Solomon 50 (NU); Underberg (-CD), McClean 598 (NH);
Coleford Nature Reserve, ‘‘Sunnyside Cottage’’, Puff 7612251 /1 (WU); Mar-
waga Mtn., Farm ‘‘Sunset’’ (-DC), Rennie 502 (NU); Bulwer-Donnybrook rd.
(-DD), Hilliard & Burtt 8036 (E, K, MO, NU, PRE, S).
228 Journal of South African Botany
——2930 (Pietermaritzburg): Weston (“‘Westtown’’) (-AA), Rehmann 7345 (K);
Elandskop, Farm ‘‘Strathavon’’, ca. 1 500 m (-BB), Ward 7645 (NU); Keerom
(-CC), Strey 10874 (NH), 10833 (NH, NU); Mooirivier distr., Johnston 578 (E),
816 (E).
—— 3029 (Kokstad): Weza, Zuurberg, nr. Transkei border (-DA), Puff 760509-1 /7
(NU, WU).
Inexact localitites: ‘‘Natal and Zululand’’, Gerrard 1338 (K, W; as ‘‘Port
Natal’’, Gerrard & M’Ken 1338 in S); ‘‘Natal’’, Cooper 2496 (E, K).
LESOTHO—2828 (Bethlehem): Leribe, ca. 1 500—1 800 m (-CC), Dieterlen
146 (K, NH, PRE, SAM); —, mountain slopes S of Mission Station, Phillips 688
(PRE, SAM); —, W slopes of Leribe Plateau, Phillips 749 (SAM).
2927 (Maseru): Mazenod (-BC), Jacot Guillarmod 799 (PRE); Morija (-DA),
Page 15964 (BOL).
2928 (Marakabei): Bokong Stores, ca. 2 350 m (-AC), Jacot Guillarmod 160
(PRE).
—2929 (Underberg): Sani Pass-Mokhotlong rd., base of Black Mtn. (-CB), Puff
761209-7/12 (WU); Sehlabathebe, 2 300—2 500 m (-CC), Jacot Guillarmod,
Getcliffe & Mzamane 315 (GRA, PRE).
CAPE—3029 (Kokstad): Kokstad, ca. 1520 m (-CB), Mogg 5190 (PRE);
Insiswa Mtn., 2 000 m (-CC), Schlechter 6450 (W, WU), Krook 2056 (W), 10595
(PRE).
——3126 (Queenstown): Stormberg, Penhoek Pass, ca. 1 830 m (-BC), Acocks
18668 (PRE, SRGH); summit of Hangklip Mtn., ca. 2 100 m (-DD), Roberts 2046) ’
(PRE). ;
——3225 (Somerset East): near Somerset East (-DA), Scott Elliot 357 (E);
Bosberg, ca. 910 m, McOwan 1414 (BOL).
——3226 (Fort Beaufort): mountain top at Oxton, ca. 1 460 m (-BA), Galpin 8360 ~
(GRA, PRE); Keiskammahoek-Hogsback rd., ca. 2 m from Hogsback village
(-DB), Puff 7609041 /1 (WU).
Uncertain localities: ‘‘Cala district’, Pegler 1793 (PRE); ‘‘Faku’s Territory’’,
Sutherland s.n. (K).
2. G. capense Thunb., Prod. Pl. Cap. 1: 30 (1794). Type: South Africa, Cape
Prov. (“‘CBS’’), Thunberg in herb. Thunberg sub no. 3308 (UPS!; NU, WU:
photos!).
G. mucronatum Thunb., Prod. Pl. Cap. 1: 30 (1794). Type: South Africa, Cape
Prov. (“‘CBS’’), Thunberg in herb. Thunberg sub no. 3342 (UPS!; NU, WU:
photos!).
G. expansum Thunb., Prod. Pl. Cap. 1: 30 (1794). Type: South Africa, Cape
Prov. (“‘CBS’’), Thunberg in herb. Thunberg sub no. 3312 (UPS!; NU, WU:
photos!).
G. capense var. expansum (Thunb.) Sond. in FI. Cap. 3: 36 (1865).
TT
—z
The Genus Galium L. (Rubiaceae) in Southern Africa 229
G. expansum var. elongatum Eckl. & Zeyh., Enum. Pl. Afr. Austr.: 370
(1836). Type: South Africa, Cape Prov., between ““Hassaquaskloof’’ and
““Breederivier’’, Ecklon & Zeyher 23268 (BOL!, SAM!; WU: photo!).
G. mucronatum var. densiflorum Eckl. & Zeyh., Enum. Pl. Afr. Austr.: 370
(1836). Type: South Africa, Cape Prov., ‘‘Vierentwintig Rivieren’’, Ecklon &
Zeyher 2327 B (GOET!, P!, SAM!, W!; WU: photo!).
G. mucronatum var. subglabrum Eckl. & Zeyh., Enum. Pl. Afr. Austr.: 370
(1836). Type: South Africa, ‘‘Tambukiland’’ (= Eastern Cape Prov.), on the
right side of the ‘‘Keyrivier’’ (= Kei R.), Ecklon & Zeyher 2327 (GOET!, P!
SAM!, W!; WU: photo!).
G. namaquense Eckl. & Zeyh., Enum. F. Afr. Austr.: 369 (1836). Type: South
Africa, Cape Prov. near “‘Heerelogement’’ (Heerenlogement), Ecklon & Zey-
her 2322 (S!, SAM!; WU: photo!).
G. garipense Sond. in Fl. Cap. 3: 37 (1865). Types: South Africa, Cape Prov.,
“‘on the Garip near Buffelvallei’’, 4 000 ft., Drege (?= Drége 7674, K!, S!,
W!); —, Natal, ‘“‘Buffaloerivier’’, Gerr. & M’K. (?= Port Natal, Gerrard &
McKen 1340 in S!; as Natal & Zululand, Gerrard 1340 in K!).
G. wittbergense Sond. in Fl. Cap. 3: 37 (1865). Type: South Africa, Cape
Prov., “‘rocky wet places in the Wittbergen’’, 6-7 000 ft. Jan. Drege (in
- herbarium Sonder).
G. capense var. minus Sond. in Fl. Cap. 3: 36 (1865). Types: South Africa,
Cape Prov. (‘‘CBS’’), Thunberg in herb. Thunberg sub no. 3342 (UPS!; NU,
WU: photos!); —, —, Wolvekop, Zeyher 773 (S!).
G. capense var. scabrum Sond. in Fl. Cap. 3: 37 (1865). Types: South Africa,
Cape Prov., Modderfonteinsberg, 4 000-5 000 ft., Drege 7682 (E!, K!, P!,
S!; WU: photo!); —, —, between Pedroskloof and ‘‘Lilly Fontein’’ (Leliefon-
tein), 3 000—4 000 ft., Drége 7683 (K!, P!; WU: photo!).
G. wittbergense var. glabrum Phillips in Ann. S. Afr. Mus. 16: 113 (1917).
Types: Lesotho, Leribe, Phillips 713 (K!, SAM!); —, —, Dieterlen 40 p.p.
(SAM!, mixed with var. wittbergense).
Perennial with a sometimes slightly woody rootstock. Stems scrambling,
sprawling or climbing, or ascending to erect, (80)150—900 mm long, with a few to
many lateral branches; up to 4 mm in diam. and + terete at the base, 0,5—3 mm in
diam. and distinctly 4-angled in the mid-stem region; glabrous or with a few
minute reversed prickles, or with + short curled whitish hairs. Longest internodes
(8)10-40 mm. Middle cauline leaves in whorls of 6-8(-10), 1-nerved, (3)5-2023)
x (0,3)0,5—2(2,5) mm, linear to narrowly lanceolate or oblanceolate, with a
sometimes brownish-white mucro at the apex; surface sometimes shiny, glabrous
or with small, forwardly directed prickles, or with short, straight hairs; margins
often reflexed, glabrous or with a few reversed prickles. Synflorescences broadly
pyramidal to + cylindrical, paracladia + many-flowered, flowers arranged in
230 Journal of South African Botany
groups of (15)12—3(1), ultimate branches with (4)3-1 small, linear bracts or
ebracteate; peduncles (1)1,5—3 mm, glabrous or scabrous, pedicels (1)1,5—4(5)
mm, + filiform, glabrous or scabrous, divaricate in fruit. Flowers 4-merous;
corolla 2-4 mm in diam., rotate, (bright) yellow, creamy yellow, greenish-yellow
or (rarely) whitish; lobes (much) longer than wide, acute or sometimes slightly
acuminate; stamina ca. %4 to % of lobe length, filaments filiform; styles % to %
fused. Fruits dry, glabrous, + tuberculate or granulate, or beset with short straight
or curled whitish hairs; mericarps subglobose, each ca. 1—2 mm wide.
Preliminary Remarks:
Sonder’s (1865) and my treatment of Galium differ considerably in regard to
the delimitation of G. capense: according to Sonder the Galium capense ‘‘com-
plex’’ consists of G. capense (with 4 varieties), G. garipense Sond. and G.
wittbergense Sond.; in the present revision, however, G. capense is split up into
three subspecies:
2A. ssp. capense
2B. ssp. namaquense (Eckl. & Zeyh.) Puff, and
2C. ssp. garipense (Sond.) Puff, with
2Ca. var. garipense and
2Cb. var. wittbergense (Sond.) Puff
G. capense var. scabrum Sond. (= G. namaquense Eckl. & Zeyh.) becomes
G. capense ssp. namaquense: although sometimes difficult to keep apart morpho-
logically from ssp. capense, it is separated geographically from the latter. G.
capense s.str. is not subdivided further: all infraspecific names used by Sonder and
other early authors are dropped because they stand for indistinguishable morpho-
logical forms. G. garipense cannot be kept apart specifically from G. capense
s.str. (particularly in the Eastern Cape Province), and therefore is treated as the
‘Eastern’ subspecies of the G. capense ‘‘complex’’. G. wittbergense, on the
other hand, merely differs from ssp. garipense in its hairiness and has a +
identical distribution range: it is, therefore, considered a variety of ssp. garipense.
2A. G. capense ssp. capense
Type: see 2. G. capense Thunb.
G. mucronatum Thunb., Prod. Pl. Cap. 1: 30 (1794). Type: South Africa,
Cape Prov. (“‘CBS’’), Thunberg in herb. Thunberg sub no. 3342 (UPS!; NU,
WU: photos!).
G. expansum Thunb., Prod. Pl. Cap. 1: 30 (1794). Type: South Africa, Cape
Prov. (‘CBS’’), Thunberg in herb. Thunberg sub no. 33/2 (UPS!; NU, WU:
photos!).
G. capense var. expansum (Thunb.) Sond. in Fl. Cap. 3: 36 (1865).
G. expansum var. elongatum Eckl. & Zeyh., Enum. Pl. Afr. Austr.: 370
(1836). Type: South Africa, Cape Prov., between ‘‘Hassaquaskloof’’ and
“Breederivier’’, Ecklon & Zeyher 23268 (BOL!, SAM!; WU: photo!).
The Genus Galium L. (Rubiaceae) in Southern Africa 231
G. mucronatum var. densiflorum Eckl. & Zeyh., Enum. Pl. Afr. Austr.: 370
(1836). Type: South Africa, Cape Prov., ““Vierentwintig Rivieren’’, Ecklon
& Zeyher 23278 (GOET!, P!, SAM!, W!; WU: photo!).
G. capense vat. minus Sond. in Fl. Cap. 3: 36 (1865). Types: South Africa,
Cape Prov. (*"CBS’’), Thunberg in herb. Thunberg sub no. 3342 (UPS!; NU,
WU: photos!);—, —, Wolvekop, Zeyher 773 (S!).
Perennial with a + woody rootstock. Stems scrambling or climbing, (80)150—
500(700) mm long, with few to many lateral branches; up to 4 mm in diam. and +
terete at the base, 0,5—1,5(2,5) mm in diam. and distinctly 4-angled in the
mid-stem region, (at least younger parts) densely covered with short, curled,
whitish hairs; often purplish. Longest internodes (8)10—35(40) mm. Middle cau-
line leaves in whorls of (6-)8(—10), 1-nerved, (3)5—12(15) x 0,5-1,5(2) mm,
linear to narrowly lanceolate or oblanceolate, with a brownish-white mucro at the
apex; surfaces often shiny, glabrous or occasionally with a few reversed prickles
on the (strongly) reflexed margins. Synflorescences broadly pyramidial to +
_ cylindrical, paracladia usually many-flowered, flowers arranged in groups of
(12)9—3(1), ultimate branches with (4)2—1 small bracts or ebracteate; peduncles
(1)1,5-3 mm, glabrous,, pedicels (1)2—4(5) mm, + filiform, glabrous,, usually
strongly divaricate in fruit. Flowers 4-merous; corolla (2)2,5-4 mm in diam.,
rotate, bright to pale yellow; lobes much longer than wide, acute or sometimes
slightly acuminate; stamina ca. half as long as the lobes, filaments thin; styles
fused to the middle. Fruits dry, glabrous, + tuberculate or granulate; mericarps :
subglobose, each ca. 1 mm wide.
Chromosome Number: Unknown. Diploid according to pollen measurements
(ch Big 2):
Habitat: In rocky grassland, in kloofs under rock overhangs, on ledges;
sometimes in moist ground near watercourses. Usually in sheltered locations. Ca.
200-1 500(1 700) m.
Flowering Period: (August) September to December (January).
Distribution (map, Fig. 14): From the South West Cape extending to the
Eastern Cape Province; also in Little, Great and Upper Karoo.
Critical Remarks: The morphological variability of ssp. capense is reflected in
the large number of synonyms. They merely refer to weakish, rather atypical
plants (G. capense var. minus) or to forms characterized by + lax, broadly
pyramidal synflorescences and long internodes (G. expansum, G. capense vat.
expansum, G. expansum var. elongatum) or + dense, narrow synflorescences and
short internodes (G. mucronatum var. densiflorum). As all these (extreme) forms
are connected to each other by a series of morphological intermediates, it does not
seem justified to uphold them taxonomically. The fairly uniform pollen diameters
of morphologically rather divergent forms (Fig. 2b)—a good indication that one
ploidy level prevails (presumably diploids; compare with pollen diameters of ssp.
232 Journal of South African Botany
G. capense
ssp. Cc.
ssp. namaquense
ssp. garipense
var. g.
var. wittbergense
var. g. & var. W.
Fic. 14.
Distribution of G. capense ssp. capense, ssp. namaquense, and ssp. garipense var.
garipense and var. wittbergense. Question marks: uncertain localities.
garipense )—are an additional argument against a further subdivision of ssp.
capense.
For occasional problems encountered in delimiting ssp. capense from ssp.
namaquense and ssp. garipense refer to the ‘‘Critical Remarks’’ section of the
latter.
COLLECTIONS
CAPE—2823 (Griekwastad): at Bloem’s Fontein (= Grootfontein) (-AC), Bur-
chell 2640 (GOET, K, P).
—3026 (Aliwal North): Eland’s Hoek nr. Aliwal North, ca. 1 400 m (-DA),
Bolus 143 (BOL).
——3124 (Hanover): Richmond, Roelofsfontein (-DA), Hanekom 1937 (PRE).
Wapadsberg pass (-DD), Maguire 715 (NBG).
——3125 (Steynsburg): nr. Middelburg (-AC), Acocks 5536 (S).
——3126 (Queenstown): Molteno distr., Looperberg, ca. 1 600 m, Mogg 2771
(PRE).
The Genus Galium L. (Rubiaceae) in Southern Africa 233
—3127 (Lady Frere): Clarksdale, Dordrecht (-AC), Taylor 5648 (NBG).
—3218 (Clanwilliam): Clanwilliam (-BB), Zeyher s.n. (K).
—3220 (Sutherland): Klein Roggeveld, Oranjefontein, ca. 1 450 m (-DC), Bond
17 (NBG); Sutherland distr., Vertaten Kloof (?spelling), Adamson D130 (PRE).
3224 (Graaff-Reinet): 25 m W of Graaff-Reinet, ca. 1 500-1 800 m (-AC),
Acocks 15865 (PRE; atypical); nr. Graaff-Reinet, ca. 700 m (-BC), Bolus 139
(BOL, PRE); between Graaff-Reinet and Mussupburg, ca. 1 500-1 800 m, Bolus
(?) 225 (BOL).
—3225 (Somerset East): Mountain Zebra National Park (-AB), Brynard 150
(PRE); Bosberg, ca. 1 450 m (-DA), Tofel (?spelling) 416 (PRE). * ” " v/ “4
—3227 (Stutterheim): ca. 7 m from Hogsback-Cathcart rd., Happy Valley, ca.
1 500 m (-AC), Johnson 1268 (GRA); banks of Toise R., ca. 820 m (-AD),
Acocks 9413 (PRE).
3318 (Cape Town): Paarlberg, ca. 300-600 m (-DB), Drége s.n. (“‘G.
expansum Thunb.”’, E, K, P, S, W); Kanon Berg (-DC), Salter 6447 (BOL, K);
Tigerberg, Nov. 38, no collector given (?Ecklon) (W).
—3319 (Worcester): Winterhoekberg, ca. 300-600 m (-AA), Drége 7680 (K,
W); Vierentwintig Rivieren, Ecklon & Zeyher 23278 (GOET, P, SAM, W);
Baviaansberg (-BA), Stokoe s.n. sub SAM 70003 (SAM), sub PRE 41989 (PRE),
ca. 1 680 m, Bond 145] (NBG); between Hottentotskloof and Karroopoort (-BA,
-BC), Pearson 4818 (B, BOL, K); Hex R. Mtns., Prospect Peak (-BC), Esterhuy-
sen 15962 (BOL, PRE); Brandvlei, ca. 360 m (-CB), Levyns 10775 (NBG);
Worcester, 300 m, Marloth 12560 (PRE); between Worcester and Villiersdorp, ca.
510 m (-CB, -CD), Bolus 5061 (BOL); Louwshoek Mtn. (-CD), Stokoe s.n. sub
SAM 59533 (SAM); Hex R. valley nr. De Doorns Bolus 13123 (BOL); Boschjes-
veld nr. Mordkuil, on the Doorn R., below 300 m, Drége 7685 (E, K, P, W),
Koo (-DB), Compton 3890 (BOL, NBG); Eendragt pass nr. Koo, ca. 1 070 m,
Levyns 7977 (BOL); Worcester distr., Andensberg, Esterhuysen 3259 (BOL),
Compton 9743 (NBG);—, Keeromsberg, Esterhuysen 9187 (BOL); Ceres distr.,
Bokkeveld Tafelberg, Esterhuysen s.n. sub BOL 31401 (BOL).
——3320 (Montagu): Pieter Meintjes, 1 150 m (-AD), Marloth 10777 (PRE);
Bantams, ca. 900 m (-BA), Compton 12141 (NBG); Witteberg S of Bantams, ca.
1370 m, Esterhuysen 30488 (BOL); Matjiesfontein, Witteberg, ca. 1070 m,
Compton 2806 (BOL, NBG); Witteberg, ca. 1 220 m, Adamson s.n. (BOL, K);
Dobbelaars Kloof, ca. 760 m (-BC), Levyns 6718 (BOL); Malagas, Esterhuysen
4451 (NBG, PRE); Bonnievale, ca. 425 m (-CC), Levyns 4589 (BOL); nr. Barry-
dale on Montagu rd., ca. 700 m (-DC), Levyns 573 (BOL).
—— 3321 (Ladismith): Between Litte Vetter R. and Garcia’s pass (-CC), Burcnell
6926 (K); W of Rooiberg pass, ca. 1070 m (-DA), Willems 1639 (NBG);
Rooiberg pass, Wurts 1611 (NBG); Swartberg, slopes below Kariegasberg, 1 300 m,
Thompson 2179 (PRE).
234 Journal of South African Botany
3322 (Oudtshoorn): Swartberg pass (-AC), Stokoe 8645 (BOL), ca. 1 030 m,
Wall 340 (S), Acocks 1429 (S); —, 11 m E from top, ca. 1 370 m, Stokoe s.n. sub
SAM 67539 (PRE, SAM); Langekloof, ca. 600—1 200 m (-CB), Ecklon & Zeyher
‘“‘100—12”’ (S); Kommassie Mtns. (‘““Komanatsie Hills’’) (-DA, -DB), Prior s.n.
(K).
3323 (Willowmore): Brandhoek, on rd. from Kouga R., ca. 460 m (-CC),
Fourcade 5092 (BOL); Joubertina (-DD), Horn s.n. sub PRE 4/964 (PRE).
—_—3324 (Steytlerville): on Kouga R., rd. from Suuranys (“‘Zuur Amys’’), ca. 275
m (-CD), Fourcade 3100 (BOL).
—3418 (Simonstown): Steenbras (-BB), Rogers 17860 (PRE); Hottentots-
Holland, Gueinzius s.n. (S); Hottentotsholland Kloof (= Sir Lowry’s Pass),
Ecklon & Zeyher 2327 (W).
—3419 (Caledon): nr. Caledon (-AB), Esterhuysen 4404 (BOL); Caledon Distr.,
Cruse 49 (S).
3421 Ree nr. Riversdale . AB), ca. 180 m, Muir 2942 | (PRE), c ca. 210 m,
Schlechter 1877 (GRA, PRE).
—3423 (Knysna): Plettenberg Bay (-AB), Pappe s.n. (K, S).
Uncertain, inexact or doubtful localities: Between ‘“Hassaquaskloof’’ and
“‘Breederivier’’, Ecklon & Zeyher 23268 (BOL, SAM); Wolvekop, Zeyher 773
(S); from Tulbaghskloof (Nieuwekloof) to Pikenierskloof, ca. 150-300 m, Drége
“140.10’’ (E); between Montagu and Triangle, Barnard s.n. sub SAM 32714
(SAM); in the Houthoek Mtns. and hills between Gauritz R. and Langekloof,
Ecklon 1836, 2326 (GOET, WU); between Swellendam and George, Mundt
(Ecklon & Zeyher) 91 (S); on the Fish R., ca. 1 370 m, MacOwan 1627 (SAM);
Stockenstrom, Scott Elliot 344 (E); between Brakrivier and Uitvlugt, ca. 900-—
1 200 m, Drége 7679 (K, P); ‘‘Kaffirland’’, Barber s.n. (S); ‘‘Regio orientalis’’
or ““CBS’’, Sparrman s.n. (S), Thunberg s.n. (P, S), Dahl s.n. (S), Prior s.n. (K),
Thorn s.n. (K), Thorn 357 (K), Scott Elliott s.n. (E), Meyer s.n. (GOET); highly
doubtful: Port Natal, Gueinzius s.n. (W); Durban, Natal, Kassner 1716 (E).
2B. ssp. namaquense (Eckl. & Zeyh.) Puff, stat. nov.
G. namaquense Eckl. & Zeyh., Enum. Fl. Afr. Austr.: 369 (1836). Type:
South Africa, Cape Prov., near ‘“‘Heerelogement’’ (Heerenlogement), Ecklon
& Zeyher 2322 (S!, SAM!; WU: photo!).
G. capense var. scabrum Sond. in Fl. Cap. 3: 37 (1865). Types: South
Africa, Cape Prov. , Modderfonteinsberg, 4 000—5 000 ft., Drége 7682 (E!,
K!, P!, S!; WU: photo!); —, —, between Pedroskloof and ‘‘Lilly Fontein’’
(Leliefontein), 3 000-4 000 ft., Drége 7683 (K!, P!; WU: photo!).
Perennial with a slightly woody rootstock. Stems scrambling or sprawling,
(150) 300-900 mm long, generally much-branched; 1-3 mm in diam., + terete at
The Genus Galium L. (Rubiaceae) in Southern Africa 235
the base, otherwise distinctly 4-angled, densely covered with very short, usually
curled whitish hairs; often purplish. Longest internodes (20)25—40 mm. Middle
cauline leaves in whorls of 6—8, 1-nerved, (4)6-8(10) x (0,5)1—2(2,5) mm, linear
to narrowly lanceolate or oblanceolate, with a brownish-white mucro at the apex;
upper and (sometimes) lower surface and margins (densely) covered with small,
forwardly directed prickles or short, straight hairs; margins often reflexed. Synflo-
rescences broadly pyramidal to cylindrical, paracladia + many-flowered, flowers
arranged in groups of (9)6—3, ultimate branches with 3-1 tiny, linear bracts;
peduncles 1,5—2,5 mm, scabrous, pedicels (1)1,5—3(4) mm, + filiform, scabrous
or more rarely subglabrous, divaricate after anthesis. Flowers 4-merous; corolla
(2)2,5—-3,5 mm in diam., rotate, yellow or greenish-yellow; lobes longer than
wide, acute; stamina short, ca. %4 to % of lobe length, filaments filiform; styles
short, fused to the middle. Fruits dry, beset with short, straight whitish hairs or
glabrous, + tuberculate or granulate; mericarps subglobose, each 1-2 mm wide.
Chromosome Number: Unknown. Diploid according to pollen measurements
(cf. Fig. 2).
Habitat: Scrambling in dense vegetation of river banks, in arid fynbos,
Renosterveld, or Erica-bush. Frequently in shady, + sheltered locations and sandy
soils. Ca. 725—1 500 m.
Flowering Period: (August, September) October to December.
Distribution (map, Fig. 14): Endemic to the Western Cape Province.
Critical Remarks: Although the morphological distinction between the poly-
morphic ssp. namaquense and ssp. capense may occasionally become rather
difficult (sometimes indumentum is the only ‘‘good’’ separation character), it was
decided to follow Ehrendorfer’s (1952, unpublished) suggestions on sheets from
the herbaria K and P to treat ‘‘namaquense’’ as a subspecies of G. capense. Its
distinct and + closed area of distribution (with hardly any overlap with ssp.
capense) supports the subspecies rank rather than varietal status (the latter would
possibly have been more appropriate if the taxon had been sufficiently scattered
geographically).
COLLECTIONS
CAPE—2917 (Springbok): Modderfonteinsberg (-DB), Drege 7682 (E, K, P,
S, W).
3018 (Kamiesberg): SW of Leliefontein (-AB), Pearson 6302 (K), between
Pedrosdorp and Leliefontein, ca. 900-1 200 m, Drége 7683 (K, P, W); between
Leliefontein and Ezelsfonteins, ca. 1 380 m (-AB, -AC), Pearson 6319 (B, BOL,
appr. ssp. capense); between Ezelsfontein and Kamiesberg, ca. 1 520 m, Adamson
1470 (BOL, PRE; appr. ssp. capense);, Kamiesbergen, Elliotsberg (-AC), Cramp-
ton 7699 (BOL).
—31 en Giftberg, ca. 760 m (-BC), Esterhuysen 22068 (BOL);
Heerenlogement (-DC), Ecklon & Zeyher 2322 (S, SAM).
236 Journal of South African Botany
3119 (Calvinia): Lokenburg, 21 m S of Nieuwoudtville, ca. 760 m (-CA),
Story 4390 (GRA, PRE); ca. 1130 m, Acocks 17374 (K, PRE).
3218 (Clanwilliam): Pakhuis pass (-BB), Salter 5042 (K), Compton 4761
(NBG), Thode A2119 (K, PRE); E-base of Piquetberg (-BC), Pillans 8641 (BOL,
PRE; appr. ssp. capense).
——3219 (Wuppertal): Cedarberg, path to Middelberg plateau, ca. 850 m (-AA,
-AC), Levyns 2227 (BOL); —, Middelberg, Bond 1327 (NBG), Esterhuysen s.n.
sub NBG 23363 (NBG), 2487 (BOL), 2517 (PRE); —. Metjiesrivier, Wagener
292 (NBG); Algeria Nature Reserve, Cedarberg, slopes of Vensterkop, ca. 730 m,
Taylor 7485 (PRE); Cedarberg Forest Reserve, NE base of Sneeuberg, ca.
900-1 200 m (-AC), Taylor 6184 (PRE).
2C. ssp. garipense (Sond.) Puff, stat. nov.
G. garipense Sond. in Fl. Cap. 3: 37 (1865). Types: South Africa, Cape
Prov., es 1e Garip near Buffelvallei’’, 4 000 ft., Drege (? = Drege 7674,
K!, S!, ae Natal, ‘‘Buffaloerivier’’, Gerr. & M’K.(? = Port Natal,
Gerrard oe Mc’ Ken 1340 in Se as Natal & Zululand, Gerrard 1340 in K!).
See “*Critical Remarks’’.
Perennial with a sometimes + woody rootstock. Stems usually ascending to
erect, (100)200—500(600) mm long, normally with + few lateral branches; up to 3
mm in diam. and + terete at the base, 0,8—2 mm in diam. and with 4 distinct,
whitish angles in the mid-stem region; glabrous, with a few minute reversed
prickles on the angles or with + short, whitish hairs. Longest internodes 10—
30(40) mm. Middle cauline leaves in whorls of 6—8, 1-nerved, (6)8—20(23) x
(0,3)0,5—1(1,5) mm, linear, often with a + distinct mucro at the apex; glabrous | or
sometimes with a few small reversed prickles on “the often +teflexéd margin. |
Synflorescences broadly pyramidal to + cylindrical, paracladia + many-flowered,
flowers arranged in groups of (15)12—3, ultimate branches with 1—2 small, linear
bracts; peduncles 1,5—2,5 mm, glabrous, pedicels (1)1,5—3(4) mm, + filiform,
glabrous, divaricate in fruit. Flowers slightly protandrous, 4-merous; corolla
2—3,5(4) mm in diam., rotate, bright yellow, sometimes creamy-yellow, yellow-
green or (rarely) whitish, lobes longer than wide, acute; stamina ca. % of lobe
length, filaments filiform; styles % to % fused. Fruits dry, + densely covered with
short, curled whitish hairs (rarely subglabrous); mericarps + globose, each 1—1,8
mm wide.
Two varieties recognized:
2Ca. var. garipense
G. mucronatum var. subglabrum Eckl. & Zeyh., Enuny Pl. Afr. Austr.: 370
(1836). Type: South Africa, ‘‘Tambukiland’’ (= Eastern Cape Prov.), on
the right side of the,.;“Keyrivier’” (= Kei R.), Ecklon & Zeyher 2327y °
(GOET!, P!, SAMI, WI; WU: photo!).
The Genus Galium L. (Rubiaceae) in Southern Africa Dy)
G. wittbergense var. glabrum Phillips in Ann. S. Afr. Mus. 16: 113 (1917).
Types: Lesotho, Leribe, Phillips 713 (K!, SAM!); —, —, Dieterlen 40 p.p.
(SAM!, mixed with var. wittbergense).
Stems glabrous or occasionally with a few minute, reversed prickles on the
angles. Pica 1 | Speiesoae el gare ana ig dohstee nelges, on nilyes or ef nodes ~
Chromosome Number: n= 11, 2n = 220 Gato nay ac nape 1 7
Habitat: In vleis, along stream banks, on steep, well-drained (moist) rocky
slopes, or in open grassland; sometimes in disturbed ground (road-sides, etc.).
Usually in sun-exposed situations or in semi-shade. Ca. (500)1 100-3 000 m.
Flowering Period: (September) October to January (February).
Distribution (map, Fig. 14): From the Transvaal, Orange Free State, Lesotho,
Natal Midlands and Uplands, Griqualand East and Transkei to the Eastern Cape
Province.
2Cb. var. wittbergense (Sond.) Puff, stat. nov.
G. wittbergense Sond. in Fl. Cap. 3: 37 (1865). Type: South Africa, Cape
Prov., “‘rocky wet places in the Wittbergen’’, 6-7 000 ft. Jan. Drége (in
herbarium Sonder; see *‘Critical Remarks’’).
Stems hairy, but without minute prickles on the angles. = </@5 <4. ful 4 dishag
Chromosome Number: n = 11, 2n = 22.
Habitat and Flowering Period: as in var. garipense.
Distribution (map, Fig. 14): From the Transvaal, South East Orange Free
State, North Lesotho, Natal Midlands and Uplands to the Eastern Cape Province.
Critical Remarks: It is somewhat uncertain which specimens are to be
considered the valid types of G. garipense: Sonder (1865) lists “‘on the Garip near
Buffelvallei, 4 000 ft., Dec. Drége’’ as type. No Galium, however, was collected
there by Drege (cf. Drege 1843, p. 50). Specimen Drége 7674 (‘‘Kraairivier,
am und im Fluss, 4 500 Fuss, Januar’’), on the other hand, is from a nearby
locality and, moreover, corresponds well with the original description of G.
garipense. It therefore seems likely that Sonder confused these two collecting sites
(on specimen Drége 7674 from the herbarium S the name *‘Galium garipense
Sond.’’ was written on the label by Sonder himself!). The second type is
“‘Buffaloerivier, Gerr. & M’K.’’: this may well refer to the specimens Gerrard &
McKen 1340 and Gerrard 1340 (in Flora Capensis, Sonder often for some
inexplicable reason omitted collection numbers). These two specimens are prob-
ably from one collection: in several herbaria collections by Gerrard & McKen are
often attributed to Gerrard only. The locality given for these identical specimens 1s
either ‘‘Port Natal’’ on one set of labels or ‘‘Natal & Zululand’’ on another. The
assumption that this collection is indeed a type is supported by the fact that the
label of the Gerrard & McKen specimen from the herbarium S was written by
Sonder himself. However, on the label he filled in ‘‘Port Natal’’ as locality and
not “‘Buffaloerivier’’ as given in Flora Capensis.
238 Journal of South African Botany
The type of G. wittbergense originates from “‘rocky, wet places in the
Wittbergen, 6-7 000 ft. Jan. Drége’’. According to Drege (1843, p. 52) this
must be collection site I a. 54 (‘‘Witbergen, an feuchten Oertern, grasreichen
Thalern und in Felsschluchten, 6 000—7 000 Fuss, Januar’’) and specimen
Drége 7676. 1 have been unable to trace this specimen. Although Sonder
mentions that it is in his own herbarium, Dr. J. Ross (personal communication)
could not find the collection at the herbarium MEL, where most of Sonder’s
herbarium is housed.
While specimens of garipense from more Eastern localities (Transvaal, Natal)
are always clearly distinct from G. capense s.str., separation of the two taxa
becomes rather troublesome where their ranges of distribution overlap (Eastern
Cape Province). For that reason alone species status cannot be maintained for
garipense. The latter is also not specifically distinct from G. wittbergense.
Differential characters for the two taxa given by Sonder (1865, p. 37: G. garipense
has “‘more diffuse habit, glabrous, often prickly stems, smaller spreading or
reflexed leaves, and longer cymes’’) are not valid: after examination of ample
herbarium material it becomes obvious that all these characters are variable. It
would therefore seem best to reduce wittbergense to varietal rank and to include it
in ssp. garipense.
G. capense ssp. garipense, like the other infraspecific taxa, is extremely
variable in habit, branching pattern, internode length and extent of the synflores-
cence. Some forms (belonging to either of the two varieties) produce glabrescent
gynoecea or fruits instead of hairy ones, and may therefore rather closely resemble
ssp. capense.
A number of specimens from West Natal’ differ from ssp. garipense (and ssp.
capense) in having very many short lateral branches with internodes shorter than
| 10 mm, needle-like (not more than 0,5 mm wide) erect to spreading leaves
arranged 1 in whorls of (8— 710 and + smaller flowers (not more than ca. 2 mm in
| diam.)., ‘As their’ position is not at present clearly understood, they have not been
given imme recognition.
COLLECTIONS
Ssp. garipense var. garipense
TRANS VAAL—2527 (Rustenburg): Bokfontein (-DB), Jenkins 7545 (PRE).
——2528 (Pretoria): Apiesrivier, 1 600 m (-AA, -AD), Rehmann 4241 (K),
Schlechter 3633 (BOL, PRE, SAM, W; = atypical); Fairy Glen, ca. 1 460 m
(-CA), Mogg 17230 (PRE, SRGH); Pretoria, Leendertz 3736 (PRE); Onderste-
poort, Theiler 9604 (PRE); Silverton (-CB), Obermeyer 146 (PRE); between
* 2929 (Underberg): Coleford Nature Reserve, ‘‘Sunnyside Cottage’’, ca. 1 500 m (-CD),
Moll 5157 (NH), Puff 761225-2/1 (WU); Polela distr., Sunset, Lot 7, Marwaga Mtn.,
ca. 2 000 m (-DC), Rennie 268 (NU).
The Genus Galium L. (Rubiaceae) in Southern Africa 239
Baviaanspoort and Vlakfontein (Mamelodi), ca. 1 370 m, Smith 1078 (PRE); Irene
(-CC), Burtt Davy 734 (PRE), Obermeyer 123 (PRE), Rogers 11416 (K), 8661
(PRE), Leendertz 668 (PRE); Rietvlei Reserve, ca. 1 520 m (-CD), Repton 3251
(PRE); 16 m SE of Pretoria on-rd. to Delmas, ca. 1 520 m, Codd 3457 (PRE).
2529 (Witbank): Witbank, ca. 1 610 m (-CC), Rogers 24302 (GRA); Middel-
burg (-CD), Jenkins 9859 (PRE); —, on the Klein Olifantrivier, Du Plessis 842
(PRE), Young A78 (PRE).
2530 (Lydenburg): Nr. Lydenburg (-AB), Wilms 574 (E, K); Belfast (-CA),
Williams 10252 (PRE); Waterval Boven (-CB), Britten 4752 (PRE).
2626 (Klerksdorp): Klerksdorp (-DC), Convent 30 (GRA).
—12627 (Potchefstroom): Witkoppen (-BB), Pharmac. Lab., Univ. Witwatersrand
4691 (K); Isaac Stegmann Nature Reserve, 1713 m, Mogg 35478 (PRE);
Potchefstroom (-CA), Louw 776 (PRE); Potchefstroom distr., Laerberg, Theron
828 (PRE).
—12628 (Johannesburg): Johannesburg (-AA), Moss 8538 (BOL), Leendertz
6190 (PRE); Modderfontein, Conrath 353 (K); Germiston, Rogers 12107 (PRE);
Heidelberg (-AD), Leendertz 7727 (PRE); Lagersport nr. Heidelberg, /Crosser
1567 (PRE); Farm ‘‘Wetter’’ (Schoongezicht 64), ca. 1 580-1 760 m (-AD?),
Mogg 24167 (SRGH); Suikerbosrand, ca. 1 680 m (-AD, -CA), Bredenkamp 990
(PRE), Repton 5358 (PRE); 18 m E of Vereeniging, Visgat, ca. 1 520 m (-CA),
Codd 4465 (PRE).
dertz 11004 (PRE).
2630 (Carolina): Carolina (-AA), Rademacher 7276 (PRE); Spitskop (-CB),
Pott 15162 (PRE); ca. 21 m E of Amersfoort on Piet Retief rd. (-CC), Puff
7701031 /1 (NU, WU); Carolina distr., Farm ‘‘Bergendal’’, ca. 1 640 m, Galpin
gs.n. sub PRE 12498 (BOL, PRE).
—2725 (Bloemhof): Leeufontein, 10 km W of Wolmaranstad (-BB), Van Wyk
729 (PRE); Boskuil, ca. 1 300 m (-BD), Sutton 238 (PRE).
—2729 (Volksrust): Volksrust (-BD), Jenkins 9315 (PRE).
—2730 (Vryheid): Farm ‘‘Oshoek’’ nr. Wakkerstroom, ca. 2 000 m (-AC),
Devenish 440 (PRE), 1238 (K, NH), Puff 770102—-1/1 (NU, WU). fe there?
Uncertain localities: Kolonie Plaats, Burtt Davy 9096 (PRE); P.P. Rust, Rogers
\ no tOrs fr
2376 (PRE). (args
ORANGE FREE STATE—2727 (Kroonstad): Heilbron, banks of Vaal R., ca.
1 450 m (-BD), Brandmiiller 140 (NBG, PRE); Kroonstad distr., nr. Vals R., ca.
1 370 m (-CB), Pont 427 (PRE); Kroonstad distr., Farm ‘‘Rondavel Noord 1475”’,
ca. 1 340 m, Scheepers 1718 (PRE). |
——2728 (Frankfort): Senekal distr., Doornkop, Goosens 641 (PRE; + atypical).
—2825 (Boshof): Krugersdrift Nature Reserve, ca. 1 380 m (-DD), Miiller 1334
(PRE).
~> Roberts-2346-(PRE;-+- atypical).
240 Journal of South African Botany
——2828 (Bethlehem): embankment of Bethlehem-Kroonstad railway line (Bethle-
hem distr.), ca. 1 500 m (-AA), Potgieter 40 (PRE: + atypical); nr. Bethlehem,
ca. 1500 m (-AB), Flanagan 2089 (PRE), Bolus : 8172 (BOL); 6 m from
Bethlehem on Kestell rd., ca. 1 700 m, Werger 25 (PRE); Arran nr. Skulpspruit, |
CA, Il 700 m, eee 105 (PRE, SRGH); -Clarens-(-€B), -Van-Hoepen_18486 -
— 4899 (aacicmith). IDARGREBERE and SEAstetn O.F.S. Botanic Gardens, ca.
1 700 m (-AC), Jacobsz 1030 (PRE), 2076 (NBG), Van der Zeyde s.n. sub PRE
41947 (NBG, PRE, SRGH); Harrismith, Sankey 82 (K); Swinburne, Rensburgskop
(-AD), Jadbasz 178 (PRE).
—— 95 (Jagersfontein): Fauresmith distr., Groenvlei, Verdoorn 1050 (PRE).
—2926 (Bloemfontein): _Bloemfontein, Ca. 1 380 m (- AA), Potts 433 (BOL),
1091 (PRE), Hanekom 831 (PRE):—, O.F. S. ieee Garden, Miiller 412 (NBG); —
Wintervalley N of Bloemfontein, Miiller 42) (PRE); Thaba Nchu Mtn. (-BB),
—3027 (Lady Grey): Zastron (-AC), Maree 92 (PRE, + atypical).
Uncertain or inexact localities: between Bloemfontein and Brandfort, Acocks
8264 (S); Kafferfontein, Kies 316 (PRE).
NATAL—2729 (Volksrust): Newcastle Distr., Farm ‘‘Boscobello’’ (-BD),
Jenkins 12468 (PRE).
2829 (Harrismith: Van Reenen, ca. | 680 m (-AD),,, Medley Wood 12110 |
(BOL, NU, SAM), Franks sub Medley Wood 12110 (NH), Bews 228 (NU); Van |
Reenen pass, ca. 1 500 m, Medley Wood 945 (E); nr. Winterton (-CC), King 1
(PRE); Farm “‘Brakfontein’’ nr. Frere, ca. 1 070 m (-DD), Acocks 10843 (NH).
—2830 (Dundee): Vant’s Drift, ca. 1 070 m (-BA), Codd 2388 (PRE).
—2929 (Underberg): summit plateau of Drakensberg nr. Giant’s Castle pass, ca.
me 000 m (-AD), Wright 1152 (E, NU, PRE; + atypical); Estcourt, ca. 1 200 m
af Xe BB), West 1829 (NH); Mooi River (-BC), Johnston 146 (E), Medley Wood 3562
(K, NH, SAM); —, banks of Mooi R. nr. Lake Hotel (-BC), Johnston 707 (E);
hills nr. Mooi River, ca. 1 220 m, Medley Wood 676 (E), s.n. (GRA); Meteor
Ridge, ca. 1 450 m, Mogg 3326 (PRE), 8087 (PRE); Kamberg Nature Reserve,
Stillerust Vlei, Puff 761218—I/1 (WU), —2/1 (WU), -1/3 (WU); Farm ‘‘Game
Pass’’ adjacent to Kamberg Nature Reserve, ca. 1 800 m, Gordon-Gray 102 (NU);
Farm “‘Allandale’’ adjacent to Kamberg Nature Reserve, ca. 1 850 m, Puff
761220-2/3 (NU, WU; + atypical), —3/1 (WU); Kamberg (-BD), Puff 761219
1/2 (NU, WU) —2/] (WU), —3/2 (NU, WU); Drakensberg Gardens (-CC), Lawson
978 (NH), 1000 (NH; + atypical); —, above Forester’s house, ca. 2 000 m (-CC),
Puff 761226—I /] (WU); nr. Bushmen’s Nek Police Post, Hilliard & Burtt 7989 (E,
K, NU, S); Thamathu pass, ca. 2 300 m, Hilliard & Burtt 8947 (NU; + atypical);
Coleford Nature Reserve (-CD), Sidey 3922 (PRE; + atypical); Mpendhle, Farm
“Tillietudlem’’, ca. 1550 m (-DB), Huntley 107 (NH, NU). Polela distr.,
“Glenariff’’, ca. 1 500 m, Rennie 658 (NU; + atypical).
The Genus Galium L. (Rubiaceae) in Southern Africa 241
2930 (Pietermaritzburg): Weston (-AA), Fisher 436 (NH, NU); Rietvlei,
Greenwich Farm (-AB), Fry 2770 (K, PRE); nr. Pietermaritzburg (-CB), Wilms
2022 (K).
—3029 (Kokstad): Weza, Ingeli slopes (-DA), Strey 6281 (NH; + atypical).
Uncertain, inexact or doubtful localities: ““Natal & Zululand’’, ‘‘Port Natal’’,
Gerrard 1340 (K), Gerrard & Mc’ Ken 1340 (S); Movelo (?Rovelo) Hills, ca. 2 100 m,
Sutherland s.n. (K); Pietermaritzburg-Drakensberg, Wahlberg s.n. (S).
LESOTHO—2828 (Bethlehem): Leribe (-CC), Dieterlen 40 (K, P, PRE); Ox
Bow Camp, ca. 2 600 m (-DC), Eubke-250-(PRE). < = /4e72<€
2927 (Maseru): Mamathes, ca. 1 700 m (-BB), Jacot Guillarmod 89 (PRE);
Monethe’s, ca. 1 700 m, Jacot Guillarmod 1918 (PRE); Tebetebeng Mill, ca.
1 500 m, Jacot Guillarmod 357 (PRE); Roma, ca. 1 700 m (-BC), Ruch 2151
(PRE); Thabana-li-Mele nr. Roma, ca. 2 600 m, Williamson 340 (K); Mafeteng,
ca. 1 700 m (-CC), Watt & Brandwyk 1924 (PRE); Thaba Tsoeu, ca. 1 700 m
(-CD), Page s.n. sub BOL 16796 (BOL); Morija (-DA), Dieterlen 1145 (PRE);
between Makhaleng and Nyakasoba, ca. 2 300 m (-DA, -DB), Jacot Guillarmod
5776 (PRE).
2928 (Marakabei): nr. Blue Mountain Pass, ca. 1 900 m (-AC), Williamson
506 (K); Mamalapi, ca. 2 750 m, Compton 21349 (NBG); Maletsunyane Falls, ca.
2 100 m (-CC), Staples 173 (PRE).
— 2929 (Underberg): Ndumeni Dome and Cleft Peak, ca. 3 000 m (-AA), Killick.
‘ 2322 (PRE, SRGH); Mokhotlong, ca. 2 400 m (-AC), Jacot Guillarmod 1162
(PRE), Liebenberg 5802 (PRE).
3028 (Matatiele): Lehaha-la-Sekhonyana, ca. 2 850 m (-AB), Jacot Guillar-
mod 183 (PRE).
Uncertain localities: Maseru distr, Ha Khotso, Songca 23 (PRE); Quthing
distr., Mphaki, ca. 2 J m, Archibald 520 (GRA); ‘‘Lesotho’’, ca. 2 200 m,
CAPE—2824 (Kimberley): Warrenton (-BB), Adams I 76 (BOL; very atypical).
——2923 (Douglas): St. Clair (-BB), Orpen 79 (NBG); on Vaal R. at confluence of
Orange R., Burchell 1729 (K).
3026 (Aliwal North): ca. 3,5 m NW of Ontspringen Station, ca. 1 400 m
(-CC), Acocks 13524 (PRE); Burghersdorp, ca. 1 500 m (-CD), Flanagan 1537
(BOL, PRE; + atypical), Guthrie 4205 (NBG; + atypical), Pocock 10810 (PRE; *
+ atypical).
3027 (Lady Grey): Dulcie’s Nek (-AD), Marais 1344 (PRE, SRGH).
3028 (Matatiele): Drakensberg, Tsatsana (Satsanna) Peak, ca. 2 850 m (-AC),
Galpin 6652 (BOL, K, PRE; atypical); 20 m W of Mount Fletcher, ca. 1 830 m
(-CC), Story 500 (PRE).
—— 3029 (Kokstad): St. Bernards (-AA), Barker 8017 (NBG; + atypical), New-
market (-AD), Krook 2699 (W; + atypical); Kokstad, ca. 1 520 m (-CB),
Coleman 317 (NH), Mogg 5051 (PRE; + atypical), Goossens 173 (PRE, +
242 Journal of South African Botany
atypical); around Kokstad, ca. 1 520 m, Tyson 1519 (BOL, K, SAM; atypical),
1991 (BOL, K, PRE, SAM; atypical); nr. Mt. Ayliff, on the Umzimhlowa R. (=
Mzimhlava), 1 500 m (-CD), Schlechter 6551 (S, W; + atypical).
——3124 (Hanover): Noupoort (-BB), Rogers 17322 (K).
—— 3125 (Steynsburg): Middelburg (-AC), Theron 203 (PRE).
——3126 (Queenstown): Jamestown, ca. 1 830 m (-BB), Gerstner 261 (PRE);
Berry Reservoir E of Queenstown (-DD), Hilner 339 (GRA, PRE; appr. var.
wittbergense).
——3128 (Umtata): Umtata, nr. Umtata R., ca. 730 m (-DB), Pegler 1602 (K; +
atypical); between Maclear and Umtata, Cherikwe Mtn., ca. 1 200 m, SE Flanagan
2646 (PRE, SAM; + atypical), Bolus s.n. (BOL; + atypical).
3226 (Fort Beaufort): Katberg, ca. 1 830 m (-DA), Shonen 4290 (CR
' Sole 408 (GRA). yen ee ee
3227 (Stutterheim): Hogsback (-CA), Rattray 4 433 “PRE): 4 oe te Kei R. nr.
Komga, ca. 460 m (-DB), Flanagan GST. (BOL, NBG, NU, SAM); King
William’s Town distr., Mount Coke, ca. 600 m, Sim 196/0 (NU, PRE).
——3326 (Grahamstown): Grahamstown (-BC), Glass 831 (NBG).
Uncertain or inexact localities: Griqualand West, on the Riet R., 1 100 m,
Marloth 906 (PRE); Griqualand East, Vaalbank, Haygarth s.n. (E, as Wood 4217
in aK; + aoe Grootontea: Cul 10 (ORD Ps Le rie plots)
f IF, 7 Se Teles Fe
COLLECTIONS A: Dregos Te 1 (see pp- 226 4)
ssp. garipense var. wittbergense
TRANSVAAL—2528 (Pretoria): Derdepoort (-CB), Leendertz 357 (PRE; + _
atypical).
2628 (Johannesburg): Suikerbosrand, Kafferskraal, ca. 1 600 m (-CA), Bre-
denkamp 86 (PRE).
Uncertain locality: Breginsel, Burtt Davy 1016 (PRE):
1 680m
(-BA), Hart 64 (NU).
NATAL—2929 (Underberg): Kamberg Nature Reserve, Stillerust Vlei (-BC),
Hilliard & Burtt 8729 (E, K, MO, NU, PRE), Puff 761218—I /3 (NU, WU); Farm
‘‘Allandale’’, adjacent to Kamberg Nature Reserve, ca. 1 900—2 000 m, Puff
7612204 /I (WU); Kamberg (-BD), Wright 2085 (NU; + atypical), Puff 761219-
1/2 (NU, WU), -2/] (NU, WU), -6/] (WU; + atypical); Highmoor Forest
Station, ca. 2 070 m (-BD), Killick & Vahrmeyer 3633 (NH); valley of Umzim-
kulu R. above Drakensberg Garden Hotel, ca. 1 680 m (-CO), Hilliard & Burtt_
7736 (E, K, MO, NU, PRE, S; + atypical); Bushman’s Nek, Thamathu pass, Ca.
2 300 m, Hillard & Burtt 8946 (NU; appr. var. garipense); Coleford Nature
Reserve, “‘Sunnyside Cottage’ (-CD), Puff 761225-2/2 (WU, + atypical); —, on
the banks of the N’gwangwane R., Puff 761227-1/1 (NU, WU).
The Genus Galium L. (Rubiaceae ) in Southern Africa 243
LESOTHO—2828 (Bethlehem): Leribe, ca. 1 500-1 800 m (-CC), Dieterlen:
40 (SAM).
CAPE—3027 (Lady Grey): Majuba Nek nr. Sterkspruit (-CB), Hepburn 148
(GRA); **Cis-Garipinia’’, on the Garip (= Oranje), between Kraai R. and
Witbergen, ca. 1 500-1 800 m (-DA?), Zeyher “118:12”" (S); Wittebergen (Wit-
berge), Ben MacDhui, ca. 2 800-3 000 m (-DB), Galpin 6651 (BOL, K, PRE).
3126 (Queenstown): Broughton nr. Molteno, ca. | 930 m (-AD), Flanagan
1586 (S, SAM); Penhoek pass (-BC), Barker 2200 (NBG); Queenstown, Hangklip
Min., ca. 2 000 m (-DD), Roberts 2024 (PRE), —, ca. 1 780 m, Galpin 1783
(PRE; appr. var. garipense). ne argu
——3225 (Somerset East): Cradock distr., Cooper 548 (BOL, E, K, W).
——3226 (Fort Beaufort): Groot Winterberg, S of Tarkastad, Farm ““Fairfield’’, ca.
1 680 m (-AD), Comins 787 (PRE).
——3228 (Butterworth): Kotana (-AA), Lussem 57 (NBG; + atypical).
3. G. monticolum Sond. in Fl. Cap. 3: 36 (1865). Type: South Africa, Cape
Prov., mountains near Cape Town, Ecklon 84 (S!; WU: photo!).
Perennial with a + woody rootstock. Stems ascending to erect, ca. 0,3—1,2 m
long, usually with many lateral branches; ca. 2,5 mm in diam., densely covered
with short, spreading hairs, distinctly 4-angled, except + terete near the® base;
sometimes purplish. Longest internodes 12—17(25) mm. Middle cauline leaves in
whorls of (6—)8—10, 1-nerved, 5—8(12) < 0,6—0,8(1) mm, linear, with a long
whitish, sometimes upturned mucro at the apex, often apparently terete due to the
strongly reflexed margins; upper surface, midrib beneath and margins densely
covered with + long, white spreading hairs. Synflorescences + cylindrical,
paracladia relatively few-flowered, flowers arranged in groups of (5)4—1, ultimate
branches with (5)3—1(0) minute, linear hairy bracts; peduncles 0,8—1,5 mm, +
thickish, hairy, pedicels 0,8-1,5(2) mm, filiform, glabrous, divaricate after
anthesis. Flowers 4-merous; corolla (1,5)1,7—2(2,3) mm in diam., rotate, colour
unknown; lobes longer than wide, acuminate; stamina ca. % of lobe length,
anthers roundish; styles short, ca. half or less fused. Young gynoeceum glabrous,
+ granulate; mature fruits not seen.
Habitat: On mountain slopes, scrambling among bushes.
Flowering Period: October to November.
Distribution (map, Fig. 15): Endemic to the Western Cape.
Critical Remarks: G. monticolum is undoubtedly a very close ally of G. capense
(ssp. namaquense and ssp. capense). The few collections available clearly differ
from the above two subspecies of G. capense by their indumentum ( a villous),
their somewhat fewer-flowered paracladia and smaller flowers and their usually
narrower leaves, but are difficult to separate otherwise (partly because G. capense
is a rather variable species producing a wide range of “‘forms’’). G. capense ssp.
namaquense is probably closest to G. monticolum, and grows in the same general
LA
<K— PRE = garpense
244 Journal of South African Botany
on ei?
var. subv. &
var. S. (=) _
var. suoglabrum var subg: pe i
a G.monticolum Z
gw G. bredasdorpense a
Gn, ec ~
ae
ieee
i Fic. 15.
Distribution of G. subvillosum var. subvillosum and var. subglabrum, G. monticolum, and
G. bredasdorpense.
area. Much more material of both is needed to see if the characteristics used here
to separate them really hold. The status of G. monticolum might then need to be
reconsidered.
COLLECTIONS
CAPE—3218 (Clanwilliam): between Witte Els Kloof and Lamberts Hoek
Berg (-BD), Pillans 9083 (BOL, PRE); slopes above “*Pickinier’s Pass’’ (Piek-
Uncertain locality: “‘Mountains near Cape Town’’, Ecklon 84 (S).
4. G. bredasdorpense Puff, sp. nov.
Herba perennis, decumbens vel suberecta; caules ca. 150-250 mm longi, ramo-
sissimi, teretiusculi basi, sursum tetragoni, dense scabriosi (saltem rami juniores),
internodiis 1—5(6) mm longis. Folia et stipulae foliaceae 6-natim verticillata,
sessisila; lamina plus minusve ovato-lanceolata, 1,5-2,5 mm longa, 0,7—1,2 mm
lata, apice mucrone albida, nitida, glabra, margine plus minusve reflexa. Cymae
axillares, 3(—1)-florae; pedunculi 0,5—1 mm longi, glabri, pedicellis 0,8-2 mm
longis, filiformibus, glabris, postfloraliter divaricatis. Corolla rotata, glabra, 1,5—2
mm diam., colore ignoto; lobi 4, plus minusve triangulares, breviter acuminati.
The Genus Galium L. (Rubiaceae) in Southern Africa 245
Ovarium glabrum, plus minusve granulatum, subglobosum. Fructus maturus non
visus.
Affinis G. capensi Thunb. sed foliis brevioribus et internodiis vrevissimis et
cymis 3(—1)-floribus axillaribus valde differt.
Perennial. Stems decumbent or suberect, ca. 150-250 mm long, with many
lateral branches; 2—2,5 mm in diam. and terete at the base, ca. 0,5—1 mm in diam.
and distinctly 4-angled in the mid-stem region; at least younger parts densely
scabrous with short, straight whitish hairs. Longest internodes 1-5(6) mm. Middle
cauline leaves in whorls of 6, 1l-nerved, 1,5-2,5 x 0,7-1,2 mm, + ovate-
lanceolate, with a + long, whitish mucro at the apex, glabrous, shiny, margin +
reflexed. Synflorescences very reduced, paracladia 3—1-flowered; peduncles 0,5—1
mm, glabrous, pedicels 0,8—2 mm, with 4(—1) leaf-like bracts, filiform, glabrous,
divaricate after anthesis. Flowers 4-merous; corolla 1,5—2 mm in diam., rotate,
glabrous, colour unknown; lobes much longer than wide, + triangular, shortly
acuminate; stamina ca. half as long as the lobes, filaments filiform, anthers +
ovate; styles ca. % fused. Gynoceum glabrous, + granulate; mature fruit not seen.
Type: South Africa, Cape Prov., Bredasdorp distr., 2 m S of Wydgelegen Post office
[= 3420 (Bredasdorp) -AD], Acocks 23175 (PRE!; WU: photo!)
Habitat: ‘‘In Kalk Coastal Fynbos’’ (notes from the type collection).
Flowering Period: November to December.
Distribution (map, Fig. 15): Endemic to the limestone formations of the
Bredasdorp district.
Critical Remarks: G. bredasdorpense seems to be a close ally of G. capense
(ssp. capense), but clearly differs in having very short internodes (frequently
shorter than the minute leaves), short, ovate-lanceolate leaves (vs. long, linear in
G. capense) and highly reduced inflorescences with leaf-like bracts and smaller
flowers.
G. bredasdorpense is known only from the type collection, and only the future
can tell whether the species is indeed strictly confined to the highly interesting
limestone area in the Bredasdorp district! which is known to be extremely rich in
endemics (Rourke, personal communication, thinks that it would be possible to list
well over 100. Exact data, however, are not available).
5. G. amatymbicum Eckl. & Zeyh., Enum. Pl. Afr. Austr.: 370 (1836). Type:
South Africa, ‘“Tambukiland’’ (= Eastern Cape Province), on the right side of
the ‘‘Keyrivier’’ (Kei R.), Ecklon & Zeyher 2328 (GOET!, S!, SAM!; WU:
photo!).
Perennial with + extensive rhizomes. Stems weak, caespitose, ca. 150-300
mm long, with many, often short lateral branches; + filiform, less than 1 mm in
1 Acocks (1975) suggests that the ‘‘Fynbos’’ of this area should be considered a distinct
veld type of its own (‘‘Kalk Coastal Fynbos’’).
246 Journal of South African Botany
diam., + 4-angled, with + long white spreading hairs or glabrous. Longest
internodes 8—10(15) mm. Middle cauline leaves in whorls of 6, 1-nerved, (2)3—
5(6) X (0,7)0,8—1 mm, linear-lanceolate to lanceolate, + acuminate at the apex;
with + long white spreading hairs on upper and lower surface or at least on
margins and midrib above and beneath. Synflorescences extremely reduced;
paracladia only 1-flowered, peduncles/pedicels (0,5)1(2) mm, + thickish, hairy or
glabrous, + arcuate in fruit. Flowers 4-merous; corolla ca. 1,5 mm in diam., +
rotate, hairy outside, whitish; lobes longer than wide, acute; stamina very short.
Fruits dry, with long + straight hairs or subglabrous; mericarps + reniform, each
ca. 1—1,5 mm wide.
Habitat: ‘‘In Acacia fields’? (Ecklon & Zeyher, 1836).
Flowering Period: December.
Distribution (map, Fig. 19): Endemic to the Eastern Cape Province.
Critical Remarks: Although known from two collections only, G. amatymbi-
cum is a ““good’’ and well marked species. It is separated from weak forms of G.
capense (presumably its closest ally) by its solitary flowers in the axils of
main-stem whorls or at the end of short lateral branches; from the remaining
Southern African taxa with highly reduced synflorescences (G. spurium ssp.
africanum, G. chloroionanthum, G. bredasdorpense) by its weak stems, minute
leaves, very short peduncles/pedicels and its indumentum.
COLLECTIONS
CAPE—3226 (Fort Beaufort): ‘‘Tambukiland’’, on the right side of the
“Keyrivier’” [= between Windvogelberg and Swartkey: Drége (1847a: 584)]
(-BB), Ecklon & Zeyher 2328 (GOET, S, SAM);—, Hogsback Mtn., Gaikas Kop
(-DB), Rattray 343 (BOL). =
6. G. bussei K. Schum. & K. Krause in Bot. Jahrb. 39: 571 (1907). Type:
Tanzania, Songea distr., Ngaka (Mgaka) valley, Busse 941 (B, EA; K: photo!).
G. stenophyllum Bak. var. flavoviride Utzschn. & Merxm. in Transact. Rhod.
Sc. Ass. 43: 57 (1951). Type: Rhodesia: Marandellas distr., Dehn 719A
(SRGH!).
G. stenophyllum auct., non Bak.
Perennial with somewhat woody rootstocks. Stems erect or semi-erect, (200)
300-600 mm long, with usually few, + short lateral branches; 1,5—2(2,5) mm in
diam., with 4 distinct, often whitish angles with short, + spreading hairs or
glabrous; nodes usually densely hairy. Longest internodes (15)25—40(55) mm.
Middle cauline leaves in whorls of 6—10, 1-nerved, (18)20-30 x 0,5—-1(1,5) mm,
linear, often apparently terete due to the strongly reflexed margins, with a distinct
brownish acumen at the apex; upper and lower surface with short, + spreading
hairs, glabrous or with minute, forwardly directed prickles above. Synflorescences
SO
The Genus Galium L. (Rubiaceae) in Southern Africa 247
+ narrowly cylindrical, paracladia + many-flowered and dense, ultimate branches
with (0)1—4 minute bracts; pedicels 1-1,5 mm, + filiform, glabrous, strongly
divaricate in fruit. Flowers slightly protandrous, 4-merous; corolla 2—2,5(3) mm in
diam., rotate, glabrous, yellow, pale yellow or greenish-yellow; lobes longer than
wide, + acuminate; stamina ca. half as long as the lobes, filaments + filiform,
anthers + elliptic; styles long, ca. % fused. Fruits dry, glabrous, + granulated;
mericarps subglobose, each 1,5—2 mm wide.
Two varieties recognized:
6a. var. bussei
G. stenophyllum Bak. var. flavoviride Utzschn. & Merxm. in Transact. Rhod.
Sc. Ass. 43: 57 (1951). Type: Rhodesia: Marandellas distr., Dehn 719A
(SRGH!).
G. stenophyllum auct., non Bak.
Stems (at least on the angles) and leaves (mainly lower surface) covered with
short, + spreading white hairs.
Habitat: In grassland (often Themeda-veld), in open Brachystegia woodland;
sometimes on rocky outcrops and streamsides. Ca. 1 200-1 800 m.
Flowering Period: October to February (March, April, May).
Distribution (maps, Figs. 16, 17): From Tanzania and Malawi to Rhodesia.
hy, Refrenes: /45°37 [ ler
6b. var. glabrum Brenan in Mem. N.Y. Bot. Gard. 8: 456 (1954). Type:
Tanzania, Mbulu distr., Ufiome Mtn., Burtt 2728 (EA, K!; NU, WU: photos!).
Stems and leaves without short, + spreading white hairs, or plants glabrous
altogether. Nodes may occasionally be + hairy.
Habitat and Flowering Period: As in var. bussei.
Distribution (maps, Figs. 16, 17): From Tanzania, East Zambia, Malawi and
Mogambique to Rhodesia.
Critical Remarks: G. bussei has consistently been confused with G. stenophyl-
lum Bak., a species not at all closely allied to G. bussei but belonging to Galium
sect. Aparinoides (Puff, 1974) and restricted to Tropical Africa. For a discussion
and clarification of this problem refer to Brenan (1953). /~
On the basis of synflorescence structure, G. bussei is subdivided into two
groups of varieties (dense, many-flowered: var. bussei, var. glabrum Brenan; lax,
diffuse, sometimes few-flowered: var. strictius Brenan, var. glabrostrictius
Brenan). Only the first group is represented in Rhodesia (the other is restricted to
Malawi, Tanzania, Zambia and Zaire), and, at least as far as Rhodesian material is
concerned, the further separation into a hairy (var. bussei) and glabrous form (var.
glabrum) appears to be worth following [Verdcourt (1976:400) considers the
present subdivision somewhat unreliable, but his investigations were based on
tropical material only]. Glabrous and hairy forms may sometimes be found
together [cf. Miller 5639 (SRGH), for example].
Journal of South African Botany
248
The Genus Galium L. (Rubiaceae) in Southern Africa 249
COLLECTIONS
var. bussei
RHODESIA—1731: Ruwa distr., Tanglewood Farm, ca. 1 500 m (-CD), Miller
5639 (SRGH), 7558 (K, LISC, P, SRGH).
1831: Marandellas dist., main road near Eagle’s Nest, ca. 1 500 m (-BA),
Davies 2969 (SRGH); —, Dehn 719A (SRGH).
1832: Inyanga distr., Inyanga, ca. 1 800 m (-BC), Boughey 465 (SRGH); —,
near village Inyanga Down, Norlindh & Weimarck 4677 (PRE, SRGH); —,
Troutbeck, Gailey 206 (SRGH); —, way to Slave Pits, Grosvenor 18 (SRGH); —,
Inyanga Falls, ca. 1 600 m, Jacobsen 3709 (SRGH); Rusape distr., Valhalla
(-CA), Dehn s.n. sub SRGH 41100 (SRGH); Umtali distr., Watsomba, Kukuransia
Irrig. Farm (-DA), Biegel 1671 (SRGH);—, Umtali Commonage (-DC), Chase 205
(SRGH).
— 1932: Melsetter distr., (-D), Williams 126 (SRGH).
Uncertain localities Makoni distr., Forest Hill, ca. 1 500 m, Eyles 723 (SAM,
SRGH); —, near Maidstone, Norlindh & Weimarck 4065 (SRGH).
COLLECTIONS
var. glabrum
RHODESIA—1730/1731: Salisbury distr., Chindamore Reserve Gomakarira,
ca. 1 500 m, Wild 3752 (LISC, PRE, SRGH).
—1731: Ruwa distr., Tanglewood Farm, ca. 1500 m (-CD), Miller 5639
(SRGH); —, Ruwa, Rockfall Farm, ca. 1 500 m, Miller 8083 (K, LISC, PRE,
SRGH).
——1832: Inyanga distr., near Inyanga (-BC), Fries Norlindh & Weimarck 2445
(SRGH); —, Juliasdals, Pienaar’s Farm, Rutherford-Smith 507 (SRGH); —, near
Darmalsay (?spelling), Whellan 713 (SRGH); —, Hopkins 8602 (SRGH); Pungwe
Hills distr., (-BD), Hopkins 7156 (PRE, SRGH); Umtali distr., Christmas Pass
(-DC), ca. 1 350 m, Matineau 296 (SRGH), “Herb. O.V.M. 7092’ (SRGH); —,
Penhalonga, ca. 1 500 m, Robinson 1857 (SRGH).
1832: Umtali distr., Umtali Commonage (-DC), Wild 461 (PRE).
— 1832/1932: Umtali distr., Matika’s Kloof, Chase 1905 (SRGH); — Engwa,
1 600 m, Exell, Mendonca & Wild 3 (LISC, SRGH).
1929/1930: Selukwe distr., Waterfall Valley, Duches 2 (LISC, SRGH); —,
Loveridge 565 (SRGH).
1930: Selukwe distr., Ferny Creek (-CA), Biegel 1495 (SRGH).
1932: Umtali distr., Himalaya Mtn., Dickers Farm (-BD), Dale SKF411
(SRGH); Melsetter distr., 15 m S of Melsetter, ca. 1 800 m (-DD), Fisher 1261
250 Journal of South African Botany
(NU, PRE, SRGH); —, Chimanimani National Park, upper Bundi, west tributary,
ca. 1 500 m Phipps 432 (PRE, SRGH); —, on Haroni R., Mavi 810 (SRGH); —,
ca. 3 m SW of Melsetter, on new Umtali rd., ca. 1 400 m, Drummond 5030
(SRGH); —, Gilbury, ca. 1 000 m (-D), Wild 3589 (LISC, SRGH); —, Williams
126 (SRGH). z
2030: Belingwe distr., Bukwa Mtn., ca. 1 300 m, (-CB), Pope, Siegel & Simon mon 1077 (fa
Mtunumashava Hill (-BB), Grosvenor 511 (SRGH). [AH
2030: Fort Victoria distr., Kyle National Park Game Reserve, near base of
‘Mtunumashava Hill (-BB), Grosvenor 51] (SRGH).
7. G. scabrelloides Puff, sp. nov. q
Herba perennis, scandens vel decumbens, caudice parum lignoso; caules
(0,3)0,5—1,0(1,2) m longi, (1)1,5—2,5(3) mm diam., tetragoni, pilis albis paten-
tibus plus minusve dense obtecti, internodiis (25)30—65(75) mm longis. Folia et
stipulae foliaceae (7)8—10-natim verticillata, basi plus minusve cuneata; lamina
linearis usque anguste lineari-lanceolata vel oblanceolata, (12)15—20(22) mm |
longa, (0,6)0,8—4(5) mm lata, apice mucrone albida, margine reflexa et grosse
retrorso-aculeata, ceterum pilis albis patentibus supra et infra obtecta, rarius solum
nervi pilis muniti. Synflorescentiae multiflorae; pedunculi 2-3 mm longi, pedi-
cellis 1—-2,5(3) mm longis, plus minusve filiformibus, pilis albis obtectis, postflora-
liter valde divaricatis. Corolla 1,8—3,5(4) mm diam., rotata, vulgo extus pilis albis ;
(sparse) obtecta, lutea vel ochroleuca vel luteo-viridis vel viridula; lobi 4, plus
minusve oOvati, acuti. Fructus subglobosus usque leviter reniformis, pilis albis
strictic patentibus obtectus, rarissime glaber; cocci (0,7)0,8—1(1,2) mm diam.
Perennial with a somewhat woody rootstock. Stems climbing or decumbent,
(0,3)0,5—1,0(1,2) m long, often with + many lateral branches; (1)1,5—2,5(3) mm
in diam., with white spreading hairs at least on the 4 distinct, often whitish angles.
“"/ Longest internodes (25)30—65(75) mm. Middle cauline leaves in whorls of
(7)8—10, 1-nerved, (12)15—20(22) x (0,6)0,8—4(5) mm, linear to linear-lanceolate
or oblanceolate, with a whitish mucro at the apex and cuneate at the base; both
surfaces, or at least midrib with white, + straight spreading hairs, t the {strongly )pes|
reflexed margins with closely set, coarse reversed prickles. Synflorescences
broadly to narrowly pyramidal, paracladia many-flowered, ultimate branches with
3—I tiny, linear to lanceolate bracts; peduncles 2-3 mm, pedicels 1—2,5(3) mm, +
filiform, hairy, strongly divaricate in fruit. Flowers slightly protandrous, 4-merous;
corolla 1,8—3,5(4) mm in diam., rotate, usually somewhat hairy outside, bright
yellow, creamy-yellow, greenish-yellow or greenish; lobes much longer than wide,
+ ovate, acute; stamina a little less than half as long as the lobes, filaments +
filiform, anthers elliptic; styles fused to the middle. Fruits dry, (densely) hairy
with straight, white spreading hairs, rarely glabrous; mericarps subglobose to
slightly reniform, each (0,7)0,8—1(1,2) mm wide; often only one mericarp de-
veloped.
The Genus Galium L. (Rubiaceae) in Southern Africa 251
Type: South Africa, Natal, Cathedral Peak Forest Reserve, Catchment -
9 200.» Puff 760314172 (WU). atchment 9, ca. 2 000
Chromosome Number: n = 11, 2n = 22.
Habitat: In forest margin vegetation, dense Leucosidea and/or Buddleia scrub,
amongs bushes along stream banks, in vleis, or (more rarely) in open grassland.
Confined to damp or wet, + shady locations. Ca. (850)1 300-2 600 m.
3
\
*y
a G. scabrelloides
G. chloroionanthum
G. bussei
var. b.
var. glabrum
var. b, & Var. ge
Fic. 17
Distribution of G. scabrelloides (whole range
only), and G. busset var. bussei and var. glabrum
), G chloroionanthum (Southern Africa
(Southern Africa only).
DS) Journal of South African Botany
Flowering Period: December to March.
Distribution (map, Fig. 17): From North West Swaziland through the South
East Transvaal, South East Orange Free State, Lesotho, Natal Midlands and
Uplands, and Transkei southward to the Eastern Cape Province.
Critical Remarks: G. scabrelloides shows close affinity to the + tropical East
African G. scabrellum but differs in having stems with much longer internodes,
longer, often broader leaves, frequently very broadly pyramidal synflorescences
with bracts at the ultimate branches, longer peduncles and often larger flowers
with acute, but not acuminate corolla lobes. Although the two species are
sometimes difficult to distinguish morphologically, species status, in my opinion,
appears most appropriate for this southern “‘vicariant’’ of G. scabrellum, particu-
larly in view of the generally narrow species concepts adopted here.
G. scabrelloides displays considerable variability in its growth form (branch-
ing pattern) and leaf size and shape, which, to a certain extent, seems to be
environment-dependent. Very narrow-leaved, weakish forms were often mistaken
for “‘G. wittbergense’’ (= G. capense ssp. garipense var. wittbergense). They
are, however, easily distinguished from the latter by the indumentum (particularly
of the gynoceum: straight hairs in G. scabrelloides; short, curled hairs in the G.
capense complex) and the leaf margins with very closely set reversed prickles.
G. scabrelloides and G. capense ssp. garipense are frequently found growing
in the same area, but usually a clear difference in ecological demands is
recognizable: G. scabrelloides prefers more sheltered, not open habitats and is
frequently found in between much higher growing vegetation. It appears to favour
somewhat wetter habitats. It was also observed that G. scabrelloides begins
flowering much later than G. capense ssp. garipense when the two taxa were seen
in the same area. I have seen no hybrids, but in my opinion it cannot be excluded
that introgression takes place between the two taxa.
Occasionally one encounters + glabrescent forms’ (subglabrous gynoeceum;
leaf surfaces with very few hairs only). Since such forms may even occur within
populations whose individuals have a perfectly normal indumentum, they of
course deserve no taxonomic recognition.
COLLECTIONS
TRANSVAAL—2628 (Johannesburg): Heidelberg Kloof, ca. 1 500 m (-AD),
Story 1613 (PRE: + atypical).
2629 (Bethal): Ermelo (-DB), Leendertz 7777 (PRE).
2630 (Carolina): Lochiel (-BB), Rogers 11478 (PRE).
2730 (Vryheid): Wakkerstroom, ca. 1 900 m (-AC), Beeton 232 (SAM);
Kastrolnek (-AD), Fitzsimons & Van Dam 26062 (PRE).
" Listed as ‘‘atypical’’ in ‘‘Collections’’.
The Genus Galium L. (Rubiaceae) in Southern Africa 253
Uncertain locality: Volksrust distr., ‘“‘Highlands’’, ca. 1 700 m, Mogg 7522
(PRE).
ORANGE FREE STATE—2828 (Bethlehem): Farm ‘“‘Dunelin’’ nr. Fouries-
burg (-CA), Potts 3087 (NBG, PRE); Farm ‘‘Dunblane’’ nr. Clarens (-CB), Potts
2984 (PRE); Golden Gate National Park (-DA), Compton 22520 (NBG), Roberts
3412 (PRE); —, rd. to Generaalskop, ca. 2 700 m, Roberts 3114 (PRE; atypical); ~
Witsieshoek (-DB), Junod 17508 (PRE); Bester’s Vlei nr. Witsieshoek, ca. 1 700 m,
Flanagan 1935 (SAM, PRE); Bester’s valley, ca. 1 900 m, Bolus 8170 (BOL);
Harrismith distr., Surit (?spelling) 1/5 (PRE).={~%’
SWAZILAND—2631 (Mbabane): hill NW of Mbabane, ca. 1 370 m (-AC),
Dlamini _s.n. (K, NBG, PRE). s« 5 PRE 4/786
NATAL—2729 (Volksrust): Farm ‘‘Glen Atholl’’ nr. Volksrust and Charles-
town, ca. 1 800 m (-BD), Smith 5630 (PRE); farm ‘‘Boscobello’’ (-DB), Jenkins
25527 (PRE).
—2730 (Vryheid): Groenvlei-Wakkerstroom rd., ca. 3-5 m from Natal/Transvaal
border (-AC), Puff 770101-3/3 (WU).
2828 (Bethlehem): Royal Natal National Park, Witsieshoek Gate, ca..2 200 m
(-DD), Hilliard & Burtt 8593 (E, K, NU); —, Mont-aux-Sources, Dordge s.n.
(NBG, P, PRE), Sim s.n. sub PRE 42019 (NU, PRE); —, Tugela valley, ca.
1 500-1 800 m, Bayer & Maclean 48 (K, PRE), Humbert 15020 (P).
2829 (Harrismith): Van Reenen pass (-AD), Kuntze s.n. (K); Umlambonja (=
Mhlambonjwa) valley, ca. 1 450 m (-CC), Schelpe 7243 (BOL), Rudatis 1576 (E,
K, S); Cathedral path, ca. 2 450 m, Schelpe 107 (NU).
——72929 (Underberg): below Organ Pipes pass, ca. 2 300 m (-AA), Edwards 1177
(NU); Giant’s Castle Game Reserve, ca. 1 500-1 700 m (-AB), Trauseld 538
(NU, PRE), Coleman 361 (NH);—, nr. Barnes shelter, Puff 761221—I/1 (NU,
WU):—, Bushman R. below hutted camp, Puff 761221-4/I (NU, WU);—, tributary
of Bushman R. on path to ‘“‘Main Cave’’, Puff 761221-3/1 (NU, WU);—, Injasuti
valley, path to “‘Battle Cave’’, Puff 760516-2/5 (WU); Cathedral Peak Forest
Reserve, Forest Research Station, catchment 3, ca. 1 850 m, Killick 1241 (K, NH,
S, SRGH);—, catchment 7, Puff 7603 14-8 /4 (WU);—, catchment 9, Puff 7603 14—
1/2 (WU),-2/2 (WU),-3/1 (NU, WU),-4// (NU, WU);—, catchment 10, Puff
761120-2/8 (WU);—, nr. Sebaaieni Cave, tributary of Ndedema, Puff 761121-3/3
(WU); Giant’s Castle, ca. 2 050-2 150 m (-AD), Symons 14579 (PRE), Bruyns-
Haylett 74 (NU); Bushman’s pass, ca. 2 200 m, West 1667 (PRE); Bushman R. nr.
Dalton Bridge (-BB), Acocks 10045 (NH); 5 m from jet. of S and N branch of
Loteni R., ca. 1 550 m (-BC), Wright 1482 (NU); Farm Allandale, adjacent to
Kamberg Nature Reserve, ca. 1 950 m, Puff 761220-4/1 (NU, WU); summit of
Kamberg (-BD), Puff 7612194 /1 (NU, WU); above Highmoor Dam, ca. 2 000 m,
Nicholson 499 (NH); nr. Bushman’s Nek police post (-CC), Hilliard & Burtt 7986
srl PRE 4/98O
254 Journal of South African Botany
(NU); Drakensberg Gardens, above forester’s house, ca. 2 000 m, Puff 761226-
1/2 (WU); Coleford Nature Reserve, ‘‘Sunnyside Cottage’ (-CD), Puff 761225—
1/2 (NU, WU); Mpendhle, ca. 1 520 m (-DB), Edwards 2518 (NU); summit of
Nhlosane Mtn. nr. Dargle, Puff 761125-3/6 (WU). WeClopaxt
——2930 (Pietermaritzburg): Nottingham Road (-ACQ), Maclean 932 (NH).
—3029 (Kokstad): Farm “‘Lynn Avis’’, 10 m from Ixopo on) Donnybrook rd., ca.
1 300 m (-BB), Crewe 51 (NH, NU); Zuurberg nr. Weza, Natal/Transkei border
(-DA), Puff 760509-1/1 to—1/5 (WU), Hilliard & Burtt 8054 (K, NU, PRE;
atypical), Schlechter 6609 (B, BOL, GRA, PRE).
Uncertain or inexact localities: N Berlin Mission station nr. Hoffenthal,
Medley Wood 3556 (BOL, K, NH, SAM); Mooirivier distr., Warley common,
Mogg 7183 (PRE); Thackleta (?spelling), Johnston 779 (E); Mohlamba ( = Dra-
kensberg?) range, ca. 1 500-1 800 m, Sutherland s.n. (K).
LESOTHO—2828 (Bethlehem): Leribe, 1 500—1 800 m (-CC), Dieterlen 33 la
(SAM, PRE); Ox Bow Camp, Tschlangana valley, ca. 2 600 m (-DC), Jacot
Guillarmod 3794 (PRE).
—2927 (Maseru): Mhlatsa’s, ca. 2 100 m (-BB), Jacot Guillarmod 464 (PRE;_
atypical); Roma, ca. 1 680 m (-BC), Ruch 1742 (PRE), 1940 (PRE), Schmitz BS
(PRE).
——2928 (Marakabei): Mountain Road, ca. 2 300 m (-BC), Jacot Guillarmod 3321
(RUH, PRE).
—2929 (Underberg): Sani Pass-Mokhotlong rd., base of Black Mtn. (-CB), Puff
761209-7/5 (NU, WU).
—3027 (Lady Grey): Liseleng valley (-BD), Coetzee 522 (PRE; atypical).
No localities given: ‘‘Basutoland’’, Cooper 2499 (K), 2500 (E, Z), Schmitz
225 (PRE). pice
CAPE—3029 (Kokstad): nr. Kokstad, ca. 1 600 m (-CB), Tyson 1422 (BOL,
PRE), 1626 (SAM, BOL; atypical); nr. Rode, ca. 1 500 m (-CC), Schlechter 6434
(B, BOL, PRE).
—3128 (Umtata): nr. Umtata, ca. 900 m (-DB), Flanagan 2858 (BOL, PRE)
f 3226 (Fort Beaufort): Katberg (-BC), Sole 408 (BOL, GRA; atypical), Schon-
land 4290 (GRA; very atypical); Auckland Forest, Amatole Mtns. (-DB), Lewis
Grant 2777 (BOL, PRE).
Happy Valley, ‘‘Fenfield’’ (-AC), Comins 1753” “GRA,” 1 BRE aypleabs- ‘Fort
Cunynghame (-AD), Sim 2853 (NU); Hogsback (-CA), Rattray s.n. sub GRA
A20321 (GRA; very atypical); Dohne Hill (-CB), Sim 1140 (PRE; atypical);
Stutterheim commonage, ca. 700 m, Acocks 9771 (PRE; + atypical); Pirie, ca.
900 m (-CC), Sim 196024 (PRE; very atypical); 3 m from Amabele, ca. 800 m
(-DA), Letries (?spelling) 53 (GRA).
——3228 (Butterworth): Nqamakwe, ca. 900 m (-AB), Rennie 399 (GRA; +
atypical).
3227 (Stutterheim): Thomas R. (-AB), Compton, 19296 (NBG; atypical),
The Genus Galium L. (Rubiaceae) in Southern Africa ASS)
Inexact or doubtful locations: Inungi Mtns. between Cedarville and Mount
435 (P; atypical).
8. G. scabrellum K. Schum. in Bot. Jahrb. 28: 113 (1899). Type: Malawi, Nyika
Plateau, Whyte (‘*Carsson’’) 269 (B, K!; NU, WU: photos!).
G. bequaertii De Wild. in Rev. Zool. Afr. 9, Suppl. Bot.: 12 (1921). Type:
Zaire, Tshilirunge, Bequaert 6047 (BR).
Perennial. Stems scrambling, ca. 450 mm long, + much-branched; ca.
1,5—2 mm in diam., with + straight, spreading hairs on the 4 distinct angles.
Longest internodes 20—25 mm. Middle cauline leaves in whorls of (6—)8(—10),
l-nerved, 10-15 x (1)1,5—2(2,5) mm, linear to narrowly elliptic, with a distinct,
whitish acumen at the apex; upper surface smooth or with a few scattered, +
straight hairs, lower surface with hairs at least on the midrib, the usually reflexed
margins with closely set reversed prickles. Synflorescences + cylindrical, paracla-
dia many-flowered, ultimate branches often ebracteate; peduncles 1-2 mm, hairy,
pedicels 1-3 mm, hairy or glabrous, divaricate in fruit. Flowers 4-merous; corolla
2-2,5 mm in diam., rotate, often hairy outside, pale yellow; lobes much longer
than wide, + triangular, acuminate; stamina ca. half as long as the lobes, filaments
+ filiform, anthers + elliptic; styles fused to the middle. Fruits dry, glabrous;
mericarps + globose, each ca. 1 mm wide.
Habitat: Shrubby grassland. Ca. 1 050 m.
Flowering Period: November.
Distribution (maps, Figs. 13, 18): Through the mountain systems of Tropical
(East) Africa (Uganda, Tanzania, Zaire, Rwanda, ? Burundi, Malawi, North East
Zambia) extending into Rhodesia.
Critical Remarks: Like G. chloroionanthum and G. bussei, G. scabrellum is a
species centred in Tropical East Africa which reaches its Southern limit of
distribution in the mountains of East Rhodesia. As already discussed, it is closely
related to G. scabrelloides, but does not seem to have any other close allies
amongst the Southern African Galia.
COLLECTIONS
RHODESIA—1832: Umtali distr., Odzani River valley (-DA, -DB), Teague
314 (BOL).
9. G. subvillosum Sond. in Fl. Cap. 3: 38 (1865). Type: South Africa, Cape
Prov., Du Toitskloof, 1 000-2 000 ft, Drege 7687 (S!, W!; NU, WU:
photos!)
Perennial with a + woody rootstock. Stems + much-branched, prostrate,
ascending or scrambling, 150—500(600) mm long, often purplish; (0,7)1—1,5 mm
in diam., with 4 + distinct, sometimes whitish angles, hairy with short, spreading
The Genus Galium L. (Rubiaceae) in Southern Africa “S57
white hairs, glabrous or nearly so. Longest internodes 10—35(40) mm. Middle
cauline leaves in whorls of 6, 1-nerved, (5)6-8(11) x (1,5)2-3,5(4,5) mm,
oblanceolate or + lanceolate, with a distinct hyaline point at the apex; with
spreading, white hairs on both surfaces and the slightly reflexed margins or
glabrous (or nearly so). Synflorescences + cylindrical, much reduced; paracladia
with 3—6 flowers arranged in groups of 3; peduncles (4)5—10 mm, thickish, hairy
or glabrous, pedicels with (0)1—2 + small, lanceolate, hairy or glabrous bracts,
(3)4—6(8) mm, slightly enlongating after anthesis, thickish to + filiform, hairy or
glabrous, divaricate in fruit. Flowers strongly protandrous, 4-merous; corolla 3—4
mm in diam., rotate, usually a little hairy outside, whitish-yellow, creamy-yellow
or purplish; lobes longer than wide, + ovate, acute; stamina nearly as long as the
corolla lobes, filaments + filiform, anthers small, elliptic; styles ca. ?/, fused, less
than 1 mm long, free ends arching downward. Fruits dry, wrinkled, with short,
straight hairs or glabrous; mericarps subglobose to + reniform, each 2—2,5(3) mm
wide; often only one mericarp developed.
Two varieties recognized:
9a. var. subvillosum
Stems and foliage, peduncles and pedicels (densely) covered with short,
spreading white hairs. Also corollas (outside) and fruits may be hairy.
Chromosome Number: n= 22, 2n = 44.
Habitat: In grassland, amongst shrubs, at the base of cliffs, on shady river
banks; always in fairly moist and cool conditions, never on dry slopes. Ca.
(450)600—1 400(1 600) m.
Flowering Period: (August) September to January.
Distribution (map, Fig. 15): Endemic to the mountains of the South West
Cape.
9b. var. subglabrum Puff, var. nov. ; ;
Caules et folia glabra vel pilis albis sparse obtecta, aliter ut in var. subvilloso.
Stems and foliage glabrous or nearly so. Also penduncles, pedicels, corolla
and fruit are usually hairless.
Type: South Africa, Cape Prov., Bains Kloof, 3 500 ft., Compton 18632 (NBG).
Chromosome Number: n = 22, 2n = 44.
Habitat and Flowering Period: As in var. subvillosum.
Distribution (map, Fig. 15): Endemic to the mountains of the South West
Cape.
Critical Remarks: The densely hairy and subglabrous varieties of G. subvillo-
sum seem to be worth separating. Both varieties have a similar range of
distribution, but var. subglabrum appears to be much less common. Although they
are sometimes found together [cf. Puff 760918—2/1 (NU, WU) and -2/2 (NU,
258 Journal of South African Botany
WU), for example], they, according to my field observations, always remain
distinct and were never connected by a series of intermediates.
Veld-burning seems to have little effect on the plants. A comparison of
individuals from an unburnt and recently burnt area (carried out at the Jonkershoek
State Forest near Stellenbosch) revealed only very insignificant differences: plants
from the burnt area showed a somewhat increased vegetative growth, slightly
longer internodes and wider leaves, but are inseparable otherwise.
The sometimes purplish colour of stems, flowers, and even leaves appears to
come about as a result of overexposure to sunlight. Purplish organs were never
observed on plants of + shady locations.
COLLECTIONS
var. subvillosum
CAPE—3318 (Cape Town): Groot Drakenstein Mtns., S slopes of Banhoek
Spitzkop, 1 220 m (-DD), Esterhuysen 11866 (BOL); Swartboskloof, Van Rens-
burg 2107 (PRE); Jonkershoek State Forest, 600-900 m, Esterhuysen 9709 (BOL,
3319 (Worcester): Ceres (- -AD), Pearson +3522 (B, BOL, K), Acocks 1879 (S):
Slanghoek Mtns., Observation Peak, SE slopes (-CA), Esterhuysen 1703 (BOL);
—, Cossacks, ca. 1 400 m, Esterhuysen 24008 (BOL); —, Slanghoek, Needle, cliffs
on § side, ca. 1 400 m, Esterhuysen 17796 (BOL, NBG); Du Toitskloof, ca.
300-600 m, Drege 7687 (S, W), Puff 760908-6/2 (NU, WU); Witteberg, ca.
1 600 m, Esterhuysen 15676 (BOL, K, PRE, SAM), Esterhuysen s.n. (WU);
Bainskloof, ca. 900—1 050 m, Schlechter 9184 (BOL, E, GRA, K, P, PRE, S, W),
Esterhuysen 25614 (BOL); Franschhoek Forest Reserve (-CC), Leighton s.n. sub
BOL 31402 (BOL); —, Berg River Hoek, ca. 900 m, Compton 13817 (NBG); —,
near Bushmen’s Castle, Salter 5736 (BOL); Wemmershoek Mtns., Tierkloof, ca.
1 200-1 500 m, Esterhuysen 17685 (BOL, PRE); Wemmershoek Peak, ledges on
W side, ca. 1 200-1 500 m, Esterhuysen 11331 (BOL, K, NBG); Drakenstein
Mtns., Duivel’s Kloof, ca. 900 m, Esterhuysen 1306 (BOL).
3320 (Montagu): Lemoenshoek Peak, below summit, ca. 1 600 m (-DD),
Esterhuysen 29491 (BOL; + atypical).
3418 (Simonstown): Sir Lowry’s Pass (-BB), Stokoe s.n. sub SAM 59534
(SAM, PRE); SE of Kogelberg, Stokoe s.n. sub SAM 59530 (BOL, SAM;-
atypical); Pringle E Peak, ca. 600 m (-BD), Esterhuysen 24660 (BOL; + atypical).
——3419 (Caledon): Dwarsberg (-AA), Stokoe 8646 (BOL, K).
COLLECTIONS
var. subglabrum
CAPE—3318 (Cape Town): Jonkershoek State Forest, head of Jonkershoek
valley (-DD), Puff 7609182 /2 (NU, WU).
3319 (Worcester): Bains Kloof, ca. 1 050 m (-CA), Compton 18632 (NBG),
1
Nee
assesses
The Genus Galium L. (Rubiaceae) in Southern Africa 259
Puff 760908—5/11 (NU, WU); Slanghoek Mtns., Witteberg, ca. 1 700 m, Es-
terhuysen 27611 (BOL);—, Slanghoek Pile, ca. 900 m, Esterhuysen 1721 (BOL):
mountains S of Wemmershoek, S side of summit, 1 450 m (-CC), Andree 794
(PRE); Franschhoek Forest Reserve, Berg River Hoek, Esterhuysen 12397 (BOL
PRE), Compton 13858 (NBG). aa eget teas ,
——3418 (Simonstown): Sir Lowry’s Pass (-BB), Puff 7609202 /4 (NU, WU);
Somerset Sneeuwkop, ca. 1 200 m, Esterhuysen 8273 (BOL, K); Hottentots-
Holland Mtns., above Lourens Ford, Esterhuysen 3512 (BOL).
3419 (Caledon): Viljoenspass (-AA), Puff 760920-3 /1 (NU, WU); SW slopes
of Victoria peak, ca. 1 0S0—1 400 m, Esterhuysen 9765 (BOL).
10. G. mucroniferum Sond. in Fl. Cap. 3: 37 (1865). Types: South Africa, Cape
Prov., Du Toitskloof, 1 000—2 000 ft., Drége 7677 (E!, K!, WU: photo!);
, near Genadenthal, 2 000-3 000 ft., Drege 7686 (E!, K!, P!, PRE!, S!;
NU, WU: photos!); no localities given: Drége 7684, 7689 (S!); in ‘‘Caf-
fraria’’, Ecklon & Zeyher s.n. (S!).
G. dregeanum Sond. in Fl. Cap. 3: 38 (1865). Type: South Africa, Cape
Prov., Du Toitskloof, Drege 7688 (E!, K!, P!, S!; NU, WU: photos!).
Perennial with a + woody rootstock. Stems with usually many short lateral
branches, scrambling, ascending or erect, 100-550 mm long; 1—2(2,5) mm in
diam., glabrous, with few reversed prickles or with white, straight spreading hairs
on the 4 distinct, often whitish angles. Longest internodes (15)20—45(55) mm.
Middle cauline leaves in whorls of 6(—8), 1-nerved, (8)10—14 x (1)1,3—2(2,5)
mm, linear-lanceolate to lanceolate, with a long hyaline point at the apex; glabrous
or with a few reversed prickles on the slightly reflexed margins, or with white
straight, spreading hairs on upper surface, midrib below and margins; often very
shiny. Synflorescences often narrowly cylindrical, much reduced; paracladia with
2-4 flowers arranged in groups of 2; peduncles 5—12(15) mm, filiform or +
thickish, glabrous or hairy, pedicels with (3)2—1(0) minute, + linear, glabrous
bracts, (4)6—10(13) mm, elongating after anthesis, + filiform, glabrous or hairy,
strongly divaricate in fruit. Flowers strongly protandrous, 4-merous; corolla
(2,5)3-4 mm in diam., rotate, sometimes with a few hairs outside, greenish-
yellow, creamy-yellow or pale green; lobes much longer than wide, acute; stamina
nearly as long as the corolla lobes, filaments + filiform, anthers small, elliptic;
styles ca. ?/, fused, less than 1 mm long. Fruits glabrous, + granulate or wrinkled,
or with short straight hairs; mericarps + globose, each (2)2,5-3 mm wide; often
only one mericarp developed.
Two varieties recognized:
b
10a. var. mucroniferum
Stems and foliage glabrous or with a few reversed prickles on the angles and
=Margins (but never with white, spreading hairs); peduncles and pedicels never
_ hairy; fruits occasionally with a few hairs.
Sf leit
ww forte
Bhésg)
260 Journal of South African Botany
Chromosome Number: n = 22, 2n = 44.
Habitat: In grassland, rocky slopes and ravines; seems to prefer + cool,
sheltered and + shady locations. Ca. (200)300—900(1 500) m.
Flowering Period: (May to July), September to December.
Distribution (map, Fig. 19): Endemic to the mountains of the South West
Cape.
10b. var. dregeanum (Sond.) Puff, stat. nov.
G. dregeanum Sond. in Fl. Cap. 3: 38 (1865). Type: South Africa, Cape
Prov., Du Toitskloof, Drege 7688 (E!, K!, P!, S!; NU, WU: photos!).
Stems (mainly angles), leaves (margins, upper surface, and midrib on lower
surface), peduncles, pedicels and fruits covered with white, straight, spreading
hairs.
Habitat and Flowering Period: As in var. mucroniferum.
Distribution (map, Fig. 19): Endemic to the mountains of the South West
Cape.
Critical Remarks: Varietal status seems most appropriate for dregeanum,
which differs from G. mucroniferum only in its hairiness. The separation character
‘more robust’ listed by Sonder (1865) only holds for the type specimen (the only
specimen known to Sonder), but is by no means consistent.
2° 23°
G. mucroniferum
a var. m. @ var,m, &
QO var. dregeanum var. d.
4 G.undulatum
@® Gamatymbicum
nie, iI),
Distribution of G. mucroniferum var. mucroniferum and var. dregeanum, G. undulatum,
and G. amatymbicum.
The Genus Galium L. (Rubiaceae) in Southern Africa 261
Var. dregeanum seems to be much rarer than var. mucroniferum.
G. mucroniferum and G. subvillosum constitute a very closely related species
pair sharing, among others, the following characters: tetraploidy, very pronounced
protandry (cf. Fig. 11), flowers with stamina nearly as long as the corolla lobes
(long, thin filaments), and reduced snyflorescences consisting of few groups of 3
flowers (G. subvillosum, Fig. 7c) or 2 flowers (G. mucroniferum, Fig. 7d) on the
paracladia. All these characters distinguish this group from the remaining Southern
African Galium species. The two species, however, are, in addition to synflores-
cence differences, easily kept apart by their leaves (short, wide: G. subvillosum
vs. long, narrow: G. mucroniferum). They may virtually grow side by side in the
field [cf. Puff 7609086 /2 (NU, WU) and -6/3 (WU)], but, according to my field
observations, do not hybridize.
COLLECTIONS
var. mucroniferum
CAPE—3318 (Cape Town): Paarl Berg (-BD), no collector given (K);
Simonsberge (-DD), Esterhuysen 8812 (BOL), Jonkershoek State Forest, Parker
, Biesievlei, ca. 300 m, Rycroft
Stokoe s.n. sub SAM 64217 (SAM).
3418 (Simonstown): W side of Sir Lowry’s Pass, just below top (-BB),
Acocks 5226 (S); near Hottentots-Hollandskloof, Ecklon & Zeyher 2327 (SAM;
not P, S).
3419 (Caledon): Houhoek Mtns., Nieuwkloof (-AA), Burchell 8035 (K);
Genadenthal, ca. 600-900 m (-BA), Drége 7686 (E, K, P, PRE, S). 1 Drege
Uncertain location: ‘‘In Caffraria’’, Ecklon & Zeyher s.n. (S).
COLLECTIONS
var. dregeanum '
CAPE—3318 (Cape Town): Stellenbosch (DD), Duthie 577 (BOL); —,
Papegaaisberg, Worsdell s.n. (K). .
3319 (Worcester): Du Toitskloof (-CA, -CC), Drége 7688 (E, K, P, S).
Highly doubtful location: *‘Port Natal’’, Gueinzius S.n. (W).
262 Journal of South African Botany
11. G. undulatum Puff, nom. nov.
G. uncinatum Lichtenst. in Bartl. & Wendl., Beitr. z. Bot. 2: 12 (1825), nom.
illeg. Type: South Africa, Cape Prov. (“‘Cap. b. Spei’’), Lichtenstein s.n.
(GOET!).
G. glabrum sensu Sond. in Fl. Cap. 3: 38 (1865), non Thunb.
Perennial with a slightly woody rootstock. Stems ascending to erect, ca.
0,3—1,5 m long, usually with few lateral branches; 1,5—2,5 mm in diam., glabrous
or with a few reversed prickles on the 4 distinct angles, often + glaucous. Longest
internodes 30— 73 9(70) mm. Middle cauline leaves in whorls of 6, 1-nerved,
(15)20-30 x (6)8— 10(11) mm, obovate to obovate-oblong, with a short hyaline
point at the apex, cuneate at the base; margins with + coarse reversed prickles,
leaves otherwise glabrous; often glaucous. Synflorescences + cylindrical, parac-
ladia 3- to + many-flowered, ultimate branches usually ebracteate; peduncles
(5)10—22 mm, glabrous, pedicels (3)5—10(20) mm, slightly elongating after
anthesis, filiform, glabrous, divaricate in fruit. Flowers 4-merous; corolla (2,5)3—4
mm in diam., rotate, whitish; lobes much longer than wide, acute; stamina very
short, much less than half as long as the lobes, filaments thickish; styles short.
Fruits dry, glabrous, granulate; mericarps + globose, each (2)2,5—3,5 mm wide.
Habitat: Among shrubs on river banks, forest margins and mountain slopes.
Always in moist to wet, + shady locations. Ca. (300)450—1 050(1 200) m.
Flowering Period: September to December (January, February).
Distribution (map, Fig. 19): Endemic to the mountains around Swellendam,
Barrydale and Heidelberg.
Critical Remarks: G. undulatum has until now been known as ‘‘G. glabrum
Thunb.’’. After an examination of the type specimen [Thunberg 3315 (UPS)],
however, it became evident that “‘G. glabrum’’ must be considered a synonym of
G. tomentosum (see G. tomentosum, ‘‘Critical Remarks’’).
The name G. uncinatum Lichtenst., considered_a synonym of ‘‘G. glabrum’’
by Sonder (1865), unfortunately is illegitimate: it is a later homonym (4 years) of
the validly published name G. uncinatum S. F. Gray (1821). The name G.
undulatum was chosen as nomen novum because this name had already been
associated (although erroneously) with G. uncinatum Lichtenst. by Steudel (1840),
who lists (p. 658) “‘G. undulatum Lichtenst. in Bartl. Wendl. sec. Ecklon’’ as a
synonym of “‘G. glabrum Thunb.’’. Ecklon’s reference to G. undulatum
Lichtenst. must have been in error for G. uncinatum, the only Galium described
by Lichtenstein.
G. undulatum is undoubtedly very closely allied to G. tomentosum with which
it shares the following characters: (usually) very wide, oblanceolate or obovate
leaves with reversed prickles on the margins (otherwise glabrous), flowers with
very short stamina, and large fruits. Of the two, G. tomentosum (dioecious,
““sex-dimorphism’’) clearly is the more derived species.
The Genus Galium L. (Rubiaceae) in Southern Africa 263
COLLECTIONS
CAPE—3320 (Montagu): Lemoenshoek Peak, slopes above Boesmansbos
Forest, ca. 1 050—1 200 m (-DD), Esterhuysen 10464 (BOL, K); Grootvaders-
bosch Forest Reserve, Boesmansbos, ca. 1 050 m, Taylor 3223 (PRE).
3420 (Bredasdorp): Swellendam (-AB), Cohen s.n. sub NBG 2340] (NBG),
Kuntze 30 (K); Swellendam Mtn., Esterhuysen 4809 (BOL); Duiwelsbosch near
Swellendam, Cruse 29 (S); Duiwelsbosch and Voormansbosch near Puspas Valley
(-AB?), Ecklon & Zeyher 2325 (PRE, S, SAM); on both sides of the Buffeljagdri-
vier, ca. 300-600 m (-BA), Zeyher 2725 (K, P, S, SAM, W); Goede Hoop Farm,
mountain slopes, ca. 600 m (-BB), Thorne s.n. sub SAM 44517 (SAM).
12. G. tomentosum Thunb., Fl. Cap. 1: 551 (1813). Type: South Africa, Cape
Prov. (‘CBS’), Thunberg in herb. Thunberg sub no. 3373 & 3374 (UPS!;
NU, WU: photos!).
G. asperum Thunb., Prod. Pl. Cap.: 30 (1794), nom. illeg, Type: South
Africa, Cape Prov. (‘‘CBS’’), Thunberg in herb. Thunberg sub no. 3302
(UPS!; NU, WU: photos!).
G. glabrum Thunb., Fl. Cap. 1: 551 (1813). Type: South Africa, Cape Prov.
(‘‘CBS”’’), Thunberg in herb. Thunberg sub no. 33/3 (UPS!; K, NU, WU:
photos!).
G. asperum var. villosum Eckl. & Zeyh., Enum. Pl. Afr. Austr.: 369 (1836).
Type: South Africa, Cape Prov., Clanwilliam distr., near Brakfontein, Ecklon
& Zeyher 2323 B (GOET!, P!, S!; WU: photo!).
G. namaquense Schlechter, nom. nud.
Rubia lanata Dinter, nom. nud.
Dioecious perennial with a woody rootstock. Stems climbing or scrambling,
ca. 0,3(0,5)—2,5(3,0) m long, with many lateral branches; up to 15(20) mm in
diam. and terete at the base, (1)1,5—3(4) mm in diam. and distinctly 4-angled in
the mid-stem region; with reversed prickles on the angles or (rarely) subglabrous,
the upper parts of the stems frequently densely villous; greyish at the base, often
purplish in the upper parts. Longest internodes (20)30-80(100) mm. Middle
cauline leaves in whorls of 6-8, 1-nerved, (12)15—25(30) x (2)3-8(10) mm,
narrowly to broadly obovate, broadly ovate, or + lanceolate, often acuminate or
acute at the apex and cuneate at the base; with densely set reversed prickles on the
often slightly reflexed margins and midrib beneath. Synflorescences extensive, ete
broadly pyramidal, paracladia many-flowered, of rather different appearance in 3
and ¢ plants. 6: peduncles (1)2—-3 mm, + thickish, subglabrous to villous, .
pedicels 1-2 mm, = filiform, hairy, strongly divaricate; ce peduncles (10)15-
30(35) mm, somewhat elongating after pollination, villous; peace are
60(90) mm after pollination, + thickish, convergent. Flowers 4-merous; ¢: corolla
(2)2,5—3,5(4) mm in diam., rotate, greenish, greenish-yellow to pale yellow,
264 Journal of South African Botany
occasionally somewhat hairy outside; lobes much longer than wide, + ovate,
acute; stamina very short, filaments thickish, shorter than the ovate anthers,
gynoeceum rudimentary, disk small. 2: corolla 3-4 mm in diam., otherwise as in
3; styles ca. 1 mm, ca. ?/, fused, disk larger than in 6; rudimentary stamina
usually recognizable. Fruits dry, granulate, glabrous or sometimes with a few
whitish hairs; mericarps + reniform, with 3 distinct furrows along the back, each
(1)1,5—2 mm wide and (2)2,5—3 mm long.
Chromosome Number: n = 22, 2n = 44.
Habitat: In semistabilized and stabilized coastal dunes, usually amongst
bushes. Inland generally scrambling in + dense scrub of dry, sun-exposed slopes
or river banks, or in dried out river beds; seems to prefer (semi-)arid areas with an
annual rain fall of usually not more than ca. 400 mm. Sea level—600 (900) m.
Flowering Period: (August) September to November (December).
Distribution (map, Fig. 20): From the Southern Namib [‘‘desert and succulent
steppe’: Giess (1971)] through Namaqualand to the South West Cape; in the
Bue Great and Mpest Karroo, and eastwards to the Eastern Cape Province.
L Gogue clud IJecF ol GYAS .
FES —
G. tomentosum
7 Fic. 20.
Distribution of G. tomentosum.
The Genus Galium L. (Rubiaceae) in Southern Africa 265
Critical Remarks: G. tomentosum and G. asperum Thunb. [an illegitimate
name: there is an earlier homonym by Schreber (1771)] had long been con-
sidered separate species (Ecklon & Zeyher, 1836; Sonder, 1865; Marloth, 1932),
since the early authors did not realize that the ‘‘two’’ species are in fact only 3 (G.
asperum *) and ¢ (G. tomentosum) plants of a single species (probably because 3
and $ plants look quite different: dioecism coupled with sex-dimorphism, see
below). Salter (1937) finally clarified the situation.
G. glabrum Thunb., described on the same page as G. tomentosum, should be
included as a synonym of the latter: it merely represents a somewhat atypical
(subglabrous), but clearly male plant of G. tomentosum.
Dioecism has evolved in a number of Galium species, but G. tomentosum is
the only Southern African species to show this phenomenon. It is also one of the
very few in which dioecism is coupled with sex-dimorphism (G. cotinoides de
Cham. & v. Schlechtend., a Chilian species not related to G. tomentosum, for
example, is another: Ehrendorfer, personal communication). Although the synflo-
rescences of d and 9 plants have an identical basic structure (cf. Fig. 7a), their
appearance is rather different: in 2 plants the densely villous peduncles and
pedicels are much elongated and + weak so that in nature they are always
pendulous (Fig. 21a, b). In 6 plants, peduncles and the much less hairy or
subglabrous pedicels are short, stiff and straight, and, as a result, the synflo-
rescences are not pendulous. At present, no explanation of the function or possible
advantage of dioecism and the long, pendulous 2 synflorescences in G. tomento-
sum can be offered, but they are certainly not an adaption to anemogamy: G.
tomentosum is clearly entomogamous, and both, ¢ and ¢ flowers produce much
nectar and were frequently found to be visited by small insects particularly in
Namaqualand.
No exact data are available about the ratio of ¢ and plants in a population,
but according to my field observations there often seemed to be a slight excess of
males, which were normally very easy to spot since they apparently always begin
flowering earlier than females. Both predominance of males and their earlier
flowering are a common and well documented phenomenon in populations of
dioecious flowering plants (cf. Godley, 1964, 1976; Lloyd, 1973). ;
The woody stems of G. tomentosum are very conspicuous and can easily reach
a diameter of ca. 15 mm near the base. The wood structure, according to Noel
(unpublished personal communication), is somewhat derived: the wood consists of
short vessel elements, narrow vascular tracheids and parenchyma. There are
however no true libriform fibres or vascular rays. There may well be a correlation
between the wood structure, unusual for the Rubiaceae, and the ecology of the
species: herbaceous, non-woody genera of various non-rubiaceous families are
known to have evolved ‘‘(semi-) desert species’ with woody stems, whose wood
structure is similarily derived (Noel, personal communication).
1G. asperum var. villosum Eckl. & Zeyher, however, refers to $ plants.
266 Journal of South African Botany
Plants of very arid habitats often appear to be deciduous: fruiting plants
without leaves were rather common amongst the herbarium material investigated,
and even in late spring I could observe in the field that numerous plants had
already dropped all their leaves with the exception of a few near the tips of the
branches.
Fic. 21.
G. tomentosum (2) in its natural habitat (Namaqualand distr., near Kamieskroon, Puff
765913 —I /5). Explanations in the text.
G. tomentosum is very variable: leaf size and shape in particular may vary
considerably (cf. Fig. 5a; note variation within one population), but branching
pattern and internode lengths also vary to a certain extent. Variation is also
noticeable in the synflorescence region: in 2 plants there is a continuous series from
forms with very many-flowered synflorescences (typical) to forms with rather few
flowers on the lateral synflorescence branches (atypical; + rare); in 6 plants the
extent of the synflorescences hardly varies. The synflorescence in females is
always villous, but not so in males: some forms are villous, others only hairy near
the tips of the synflorescence branches, and some (rather infrequently) are almost
glabrous (such forms have occasionally been confused with G. undulatum). None
of these variable characters are, however, correlated so that it does not seem
appropriate to subdivide the species further.
The Genus Galium L. (Rubiaceae) in Southern Africa 267
COLLECTIONS
ORANGE FREE STATE—2925 (Jagersfontein): Brakfontein (-DA), Zeyher 50
(S).
CAPE—2816 (Oranjemund): Koeboes-Grootderm rd., ca. 7-10 m from Koe-
boes (-BD), Puff 760911-1 /1 (WU).
—2817 (Vioolsdrif): Koeboes (Kuboos) (-AC), Pillans 5402 (BOL); Stinkfontein
(-CD), Pearson 5527 (B).
—2917 (Springbok): Springbok-Kleinsee rd., ca. 1 m from turn-off to Komaggas,
nr. Buffelsrivier (-DA), Puff 760912-1/1 (NU, WU); Spektakelbergpas, ca. 8 m
from Springbok, Puff 760912-6/4 (NU, WU); nr. Spektakel, ca. 250 m, Bolus
9420 (BOL); hills at Okiep (-DB), Pillans 4971 (BOL); between Nababiep and
Modderfontein, ca. 900 m, Bolus 9419 (BOL); hills on W side of Springbok, ca.
980 m, Acocks 16488 (PRE); ca. 5 m from Springbok on Pofadder rd., Puff
760910-2/1 (NU, WU); Hester Malan Flower Reserve nr. Springbok, Puff
760910-1 /1 (NU, WU); Droedap (-DD), Esterhuysen s.n. sub PRE 41998 (PRE).
—2918 (Gamoep): Brak R., ca. 500 m (-A, -C), Pearson 3899 (B, BOL).
——2922 (Prieska): around Prieska (-DA), Bryant 227 (K, PRE).
2923 (Douglas): St. Clair, banks of Vaal R. (-BB), Orpen 5967 (BOL); St.
Clair-Douglas, Orpen 163 (SAM).
—3017 (Hondeklipbaai): margin of dry Spoeg R. bed (-AD), Pillans 18034
(BOL); Ara Koop, ca. 700 m (-BB), Schlechter 11236 (B, E, GRA, K, S, W);
Kamieskroon, Thorne s.n. sub SAM 48889 (SAM), sub NBG 23394 (NBG); nr.
Kamieskroon, on rd. to Leliefontein, Puff 760913-1/5 (NU, WU); Bowesdorp,
Stokoe 8214 (BOL, PRE, SAM); S of Brakdam (-BD), Pearson 5627 (B, BOL,
K).
—3018 (Kamiesberg): 3 m E of Kamieskroon, W slopes of Kamiesberg, ca.
1 100 m (-AC), Acocks 14988 (PRE); nr. Garies, foothills of Kamiesbergen (-CA),
Esterhuysen 1409 (NBG, PRE); Wolfhok-Garies rd., ca. 5 m from Garies, Puff
760913-2/12 (NU, WU).
—3019 (Loeriesfontein): Loeriesfontein Ravine, ca. 800 m (-CD), Pearson
4843 (BOL, K).
3118 (Vanrhynsdorp): Ebenezar, on the Olifants R., below 300 m (-CA),
Drége 7671 (K, W); Nardouw (-DC), Maguire 1048 (BOL, NBG).
3119 (Calvinia): Bokkeveld, Nieuwoudtville, 700 m (-AC), Marloth 7802
(PRE); Botterkloof pass (-CD), Hall 3879 (NBG).
—3218 (Clanwilliam): Verlorevlei (-AD), Compton 15064 (NBG); Graafwater
(-BA), Zinn 64219 (SAM); nr. Piketberg (-DA, -DD), Bolus 14284 (NH), Gulline
2617 (NBG): nr. Brakfontein, Ecklon & Zeyher 23238 (GOET, P, S, WU),
Drége ‘‘76.1’’ (GOET).
268 Journal of South African Botany
—3219 (Wuppertal): between Citrusdal and The Baths (-CA), Puff 760913-1/1
(NU, WU).
—_3220 (Sutherland): Houthoek (-CA), Hanekom 745 (PRE).
——3222 (Beaufort West): Nieuwefeld (-BA), Drége s.n. (S); between Beaufort
and Renosterkop, ca. 760-910 m (-BB, -BC), Drege s.n. (K); Sunnyside
(Beaufort West distr.), Esterhuysen 2729 (PRE).
—3224 (Graaff-Reinet): nr. Graaf-Reinet, ca. 760 m (-BC), Bolus 144 (BOL,
K), 145 (BOL).
3225 (Somerset East): Baroda, 15 m N of Cradock, ca. 950 m (-BA), Dyer
1044a (GRA, PRE); Somerset East (-DA), Bowker s.n. (K); Somerset East distr.
(?): between Zuurbergen and Klein-Bruitjeshoogte, Drége s.n. (‘‘tomentosum a’’,
E, K, P, S, W).
——3318 (Cape Town): In fields nr. Hopefield (-AB), Letty 134 (PRE); Darling,
ca. 75 m (-AD), Gulline 2104 (NBG); nr. Darling, ca. 110 m, Bolus 12702 (BOL,
NH, PRE), Puff 760915-—I/1 (NU, WU); Groot Post, Salter 6471 (BOL); Contre-
berg, Bachmann 1100 (K, P); around Mamre (‘‘Groenekloof’’) (-CB), Zeyher s.n.
sub SAM 10675, 37551 (SAM); Mamre Hill, Compton 9829 (NBG); Blaauwberg
(Blouberg), Stokoe 67538 (SAM); Blaauwbergstrand (-CD), Maguire 1279
(NBG); Milnerton (-CD, -DC), Andrae 389, 392 (PRE); Harmonie (-DC), Michell
238 (BOL, PRE); mountainous places at Klapmuts, ca. 150-460 m (-DD), Ecklon
& Zeyher s.n. sub PRE 20106 (PRE; as “‘Drége 92.10’’ in GOET, WU).
—3319 (Worcester): nr. Karroo Poort, Swartruggens Mtns. (-BA), Stokoe 7537
(SAM); ‘“Veld Reserve’’ nr. Worcester (-CB), Olivier 152 (PRE), Van Breda 36
(PRE); Hex River nr. De Doorn, ca. 500 m (-CD), Bolus 11899 (PRE); entrance to
Hex R. valley, Davidson 45 (SAM).
——3320 (Montagu): Snyders Kloof (-AB), Foley 139 (PRE); Matjiesfontein, ca.
900 m (-BA), Compton 2715 (BOL); —, (‘‘Maggiesfontein’’), Rehmann 2939
(K); environment of Matjiesfontein, Witteberg at Whitehil, Humbert 9750 (P);
foot of Witteberg (-A, -B), Compton 2528 (BOL); Witteberg Kloof, Adamson s.n.
(BOL); Ngaap Kop, ca. 1 070 m, Compton 12620 (NBG); Dobbelaars Kloof, ca.
700 m (-BC), Levyns 6663 (BOL; atypical); Montagu baths (-CC), Page s.n. sub
PRE 42011 (PRE); Baden Kloof, ca. 500 m (-CC?), Compton 18364 (NBG), Lewis
s.n. sub SAM 59532 (SAM), Levyns 7941 (BOL); Kochmannskloof (Kogmans
Kloof) (-CC), Zeyher s.n. (SAM).
——3321 (Ladismith): Bergkloof, ca. 7 m from Herbertsdale (-DD), Puff 760922—
1/1 (NU,WU).
—3322 (Oudtshoorn): De Rust, Farm ‘‘Doornkraal’’, 400 m (-BC), Dahlstrand
1575 (GRA), 2183 (PRE), 2454 (PRE); George (-CD), Ecklon & Zeyher “94.12”
(S.).
—3323 (Willowmore): Willowmore distr., Wolvefontein, Anderson s.n. sub
SAM 3755 (SAM).
The Genus Galium L. (Rubiaceae) in Southern Africa 269
——3324 (Steytlerville): Kouga R. Poort, ca. 300 m (-DA), Fourcade 3083
(BOL, K); on the Zwartkoprivier (-DB), Zeyher 225 (K, S, SAM), 2720 (S).
3325 (Port Elizabeth): Dunbrodie, Sundays R. valley, ca. 60-120 m (-BC),
Rennie 406 (BOL, atypical); Uitenhage (-CD), Ecklon & Zeyher ‘‘2-I’’ (S);
Redhouse (-DC), Paterson 96 (GRA), 96a (GRA, BOL); Uitenhage-Coega rd., ca.
1,3 m from Coega, Puff 760924-1/1 (NU, WU).
—3326 (Grahamstown): Alicedale (-AC), Cruden 66 (GRA); Cradock rd.,
6-7 m outside Grahamstown (-BC), Stauffer 5191 (K, P, PRE), 5/92 (P, PRE);
Alexandria-Pt. Alfred rd., ca. 5 m from Alexandria (-CB), Puff 760924-2/1 (WU);
Pt. Alfred (-DB), Marloth 10201 (PRE); Albany distr., Albany, Bowker s.n. (K).
—3418 (Simonstown): Rondevlei (-AB), Compton 24362 (NBG); nr. Muizen-
berg, 6 m, Schlechter 1487 (B); Fish Hoek Bay, ca. 15 m, Bolus 4834 (BOL);
Kommetjie, Brain s.n. (SRGH); Witsand Bay, Salter 6386 A, B, C (BOL, K, S);
Southeast Point, Wolley Dod 2991 (K); Cape of Good Hope Nature Reserve, nr.
Cape of Good Hope (-AD), Puff 760917-I/1 (WU); Swartklip (-BA), Puff
760919-2/14 (NU, WU), Esterhuysen 26456 (BOL); nr. Zeekoe Vlei (-BA,
-BB), Wolley Dod 872 (BOL); Faure (-BB), Parker 4509 (BOL, K, NBG);
Hottentots-Holland Mtns., Zeyher s.n. (S), Liuvenberg (?spelling) s.n. (S), no
collector given (K); surroundings of Somerset West, in Hottentotsholland, Ecklon
& Zeyher ‘‘83’’ (S); E of Cape Hangklip (-BD), Rourke 704 (NBG, S).
3419 (Caledon): Bosjesveld (-AA), Van Zijl s.n. (SAM); Friekkie se Baai
(-AC), Leighton 1556 (BOL), E of Betty’s Bay, Puff 760920-1/1 (NU, WU);
between Elim and Gansbaai (-CB, -DA), Stokoe s.n. sub SAM 64218 (SAM).
—3420 (Bredasdorp): ca. 1,5 m outside Stormsvlei on Stormsvlei-Bonnievale
td. (-AA), Puff 760921-1/1 (NU, WU); Swellendam (-AB), Ecklon & Zeyher
**98_9°° (S): nr. Swellendam, Zeyher s.n. sub SAM 16075 (SAM); nr. Bontebok
(-AB, -BA), Wall s.n. (S); Struis Bay (-CA, -CC), Esterhuysen 4433 (BOL),
Hafstrém & Acocks 2242 (PRE, S).
3421 (Riversdale): Stilbaai (-AD), Muir 120 (GRA), 5101 (BOL, PRE).
Uncertain or doubtful localities: Little Namaqualand, Waterkloof at Doorn-
poort, Pillans 5370 (BOL); “Stellenbosch, Worcester, Swellendam and Uiten-
hage’’, Ecklon & Zeyher 2323 (GOET, P, S, SAM, W); ‘“Karroo’’, Ecklon &
Zeyher 2324 (GOET, P, S, SAM, W); ‘‘Vumansberg”’ (?spelling), Julinek s.n.
(W); “Regio orientalis’? or ‘“CBS’’, Sparrman s.n. (S), Thunberg s.n. (S, 4
sheets), Wolley Dod 3680 (BOL), Drége s.n. (K), Zeyher s.n. (K), Zeyher herb.
no. 347 (P), Dahl s.n. (S), Bauer 94 (W), Scholl s.n. (W).
13. G. chloroionanthum K. Schum. in Bot. Jahrb. 30: 417 (1901). Type:
Tanzania, Rungwe Mtn., Goerze 1162 (B, BM, K!; NU, WU: photos').
Perennial. Stems climbing or creeping, up to 2,3 m long, with few te beta
branches: 1,5—2 mm in diam. (mid-stem), with + large, hyaline reversed prickles
on the 4 distinct, whitish angles, and often densely hairy at the nodes. Longest
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The Genus Galium L. (Rubiaceae) in Southern Africa ial
internodes 55-100 mm. Middle cauline leaves in whorls of 6, 1-nerved, (22)25—
35(40) x (5)7—12(15) mm, (broadly) elliptic to obovate, abruptly acuminate at the
apex, cuneate at the base; upper surface glabrous or with a few short, straight
hairs, margins and midrib (beneath) with coarse, + hyaline reversed prickles.
Synflorescences extremely reduced; paracladia 1-flowered, peduncles 0,4—0,8 mm
before, 10-20 mm after anthesis, pedicels 0,5—1 mm, with 1(2) + large to minute
bract(s), glabrous. Flowers 4-merous; corolla ca. 3—4 mm in diam., rotate, with
a few hairs outside, greenish-white to greenish-yellow; lobes about as long as
wide, + acute; stamina short. Fruits dry, densely covered with brownish, flattened
(?), hooked hairs; mericarps + reniform, each 1—1,5 mm wide.
Habitat: In forests. Ca. 1 800—2 000 m.
Flowering Period: December to February.
Distribution (maps, Figs. 17, 22): In tropical East Africa from South Sudan
(according to Ehrendorfer & Verdcourt, 1973, and Verdcourt, 1976; no specimens
seen by me and, therefore, not shown in map, Fig. 22), Uganda, East Zaire,
Rwanda, Burundi, Kenya to Tanzania, and southward to South Malawi and
Rhodesia.
Critical Remarks: The closest allies of the + tropical G. chloroionanthum,
which just reaches its southern limit of distribution in the mountains of East
Rhodesia, seem to be G. brenanii Ehrend. & Verdc., G. ruwenzoriense (Cortesi)
Chiov., and G. aparinoides Forssk., all of which are centred in Uganda, Kenya
and Tanzania (Ehrendorfer & Verdcourt, 1973). It does not seem to have close
affinities to any Southern African species.
COLLECTIONS
RHODESIA—1832: Inyanga distr., Rhodes Inyanga National Park, Pungwe
Gorge, ca. 1 800 m (-BD), Chase 5664 (LISC, PRE, SRGH).
1932: Umtali distr., Himalaya Mtn., Banti North, ca. 2 000 m (-BD), Wild
4517 (LISC, K, SRGH).
142 G. spurium L. ssp. africanum Verdc. in Kew Bull. 30: 324 (1975). Type:
Kenya, Kiambu distr., Muguga, Milne-Redhead & Taylor 7147 (K!; NU, WU:
photos!).
G. horridum sensu Eckl. & Zeyh.: Enum. Pl. Afr. Austr.: 370 (1836), non
Thunb.
G. aparine sensu Sond. in Fl. Cap. 3: 38 (1865), non L.
Annual. Stems weak, prostrate to semi-erect, (0,12)0,3-ca.2 m long,
few lateral branches; (0,8)1,5—2,5 mm in diam., with reversed prickles on the 4
distinct angles. Longest internodes (20)35-75 mm. Middle cauline leaves in
whorls of 6-8, 1-nerved, (8)15—38(45) x (1,5)3-6(7) mm, linear-lanceolate to
obovate, with a distinct, + filiform acumen at the apex, cuneate at the base; upper
with +
272 Journal of South African Botany
surface glabrous or with a few scattered + straight or curled hairs, midrib
(beneath) and margins with coarse reversed prickles. Synflorescences extremely
reduced; paracladia 1(2)-flowered, peduncles/pedicels (8)12—20(35) mm when in
fruit, + arcuate, with reversed prickles or (seldom) glabrous; pedicels occasionally
separated from peduncle by a small, + linear bract (in that case paracladia often
2-flowered). Flowers 4-merous; corolla (1)1,2—1,8(2) mm in diam., rotate, gla-
brous, greenish, greenish-white or creamy-white; lobes longer than wide, + ovate
to triangular, pointed; stamina very short, filaments + thickish, anthers roundish;
styles short. Fruits dry, with non-tuberculate, white hooked hairs less than 0,5 mm
= long; mericarps + globose, each (1,8)2,5—4 mm wide.
Chromosome Number: n = 20, 2n = 40.
Habitat: Amongst shrubs along river banks, in forest margin vegetation, scrub
forests, rocky mountain slopes, or (seldom) in dune forest margins; occasionally in
cultivated ground, disturbed areas and margins of plantations. Seems to prefer
sheltered, damp and + shady locations. Ca. 20—1950 m.
Flowering Period: All year round.
Distribution (maps, Figs. 23, 24): From Ethiopia (Cufodontis, 1965, as G.
aparinoides Forssk.), Somali Republic, Socotra (not shown in map, Fig. 23), East
and West Sudan, Zaire, Cameroun: Cameroun, } Mtn., Uganda, enya, Rwanda,
Burundi, Tanzania and Malawi to South Africa. “According to Verdcourt (1976)
also in Rhodesia (no specimens seen by me and, therefore, not shown in map, fig.
23).
Critical Remarks: The identity of the plant long identified as G. aparine L. in
Southern (and Eastern) Africa and often thought to be an introduction from Europe
(Sonder, 1865; Adamson & Salter, 1950) is now firmly established: it must be
included in G. spurium L. as proposed by Verdcourt (1975), and is best treated as
a subspecies of its own. One of the main reasons for this conclusion is the plant’s
chromosome number: G. spurium L. has a chromosome base number of x = 10
(ssp. spurium is diploid and has n = 10, ssp. africanum is tetraploid and has n =
20), whereas G. aparine L. has a base number of x = 11 (cf. Puff, 1977).
While plants from the Transvaal and Natal always agree perfectly well with
the description and type of G. spurium ssp. africanum, the situation seems
somewhat more problematical in the Cape Province: in addition to ‘“‘typical’’
collections, there are a number of specimens that differ in having somewhat more
extensive synflorescences (+ long paracladia with three to four, and even six
flowers). Such forms bear a close morphological resemblance to both G. spurium
ssp. spurium and G. aparine. One is tempted to conclude that they are
introductions which, in the Cape Province, occur together with the indigenous G.
spurium ssp. africanum. No chromosome counts of these critical (and rare) plants
are available, and, unfortunately, pollen measurements (Fig. 3d) do not contribute
much to the solution of the problem either: average pollen diameters are somewhat
smaller than those of G. spurium ssp. africanum, but yet there is a certain amount
274 Journal of South African Botany
G spurium
ssp, africanum
G. spurium - aparine
complex
Fic. 24.
Distribution of G. spurium ssp. africanum in Southern Africa; distribution of plants
belonging to the ‘‘G. spurium-aparine complex’’ (explanations in the text).
of overlap. Pollen grain size would fit into the range given for the diploid G.
spurium ssp. spurium (cf. Moore, 1975), but on the other hand, the collections in
my opinion more closely resemble G. aparine, which, however, is polyploid.
A detailed study (which must include karyological investigations), is therefore |
needed to solve the problem. These critical collections at present can only
provisionally be identified as ““G. spurium-aparine complex’’.
COLLECTIONS
TRANSVAAL—2329 (Pietersburg): Houtbosh (-DD), Rehmann 6031 (K).
—2630 (Carolina): Maviristad (-CA), Pott 4886 (BOL), 15152 (PRE).
2730 (Vryheid): Farm ‘‘Oshoek’’ nr. Wakkerstroom, ca. 1 900 m (-AC),
Devenish 807 (PRE), Puff 770102-3 /1 (NU, WU).
NATAL—2929 (Underberg): Kamberg Nature Reserve, Gladstone’s Nose
ridge, ca. 1 830 m (-BC), Wright 837 (NU), Puff 760418-1/6 (NU, WU); Farm
“Game Pass’’, Gladstone’s Nose Ridge, adjacent to Kamberg Nature Reserve, ca.
The Genus Galium L. (Rubiaceae) in Southern Africa Dis)
1 980 m, Puff 760418—1/I (NU, WU), -//3 (NU, WU); Bulwer-Impendhle rd., on
Ngelotsontwana stream (-DA), Hilliard & Burtt 7610 (E, K, NU); Polela, ca.
1 200—1 500 m, Medley Wood 4585 (K, NH).
2930 (Pietermaritzburg): Curry’s Post (-AC), Pickworth s.n. (NU); Lion
River-Dargle rd., roadside nr. Dargle (-CA), Puff 761125-1 /1 (WU).
3029 (Kokstad): Zuurberg nr. Weza, ca. 1 300 m (-DA), Schlechter 6604 (B,
BOL, PRE).
CAPE—3018 (Kamiesberg): Leliefontein, ca. 1 550 m (-AB), Levyns 4066
(BOL).
——3119 (Calvinia): nr. Calvinia, on rd. to Clanwilliam, ca. 900 m (-BD), Levyns
5070 (BOL); Calvinia distr., Ekerdam, Taylor 2689 (NBG).
——3126 (Queenstown): Queenstown, ca. 1 900 m (-DD), Galpin 1800 (GRA).
——3219 (Wuppertal): Agtertuin, ca. 240 m (-AA), Schlechter 10862 (E, GRA, K,
P, PRE, S, W).
3220 (Sutherland): Klipbanksrivier, Roggeveld foothills, ca. 900 m (-CB),
Acocks 16950 (PRE).
3226 (Fort Beaufort): jct. Queenstown valley and Zwartkei, ca. 1 200-1 350 m
(-BB), Galpin 8125 (PRE).
—3227 (Stutterheim): King William’s Town, ca. 600 m (-CD), Sim 1 9606 (PRE,
WU).
——3228 (Butterworth): Idutywa distr., Pegler 734 (PRE).
3318 (Cape Town): summit of Contreberg (-AD), Pillans 6908 (BOL);
Kirstenbosch, Nursery Gorge, Table Mtn. (-CD), Esterhuysen 352 (BOL); Paarl-
berg (-DB), Drége 7670 (E, K, P, PRE).
3319 (Worcester): Hex River Mtns., E slopes of Prospect Peak, ca. 900 m
(-BC), Esterhuysen 15937 (BOL, PRE); Klein Drakenstein Mtns., Kasteelkloof
Catchment, ca. 680 m, Kruger 1450 (PRE).
3326 (Grahamstown): Alicedale (-AC), Cruden 67 (GRA); Port Alfred (Kowie
, East) (-DB), Britten 764 (GRA, PRE), Dyson s.n., Nov. 1915 (PRE), June 1916,
~_Dyson 89 (GRA).
3418 (Simonstown): Orange Kloof (-AB), Salter 7822 (BOL); Hout Bay Golf
Course, Acocks 676 (PRE); Simon’s Bay, Wright s.n. (K).
3424 (Skoenmanskop): nr. Sea View, margin of dune forest, ca. 60 m, Acocks
21429 (PRE). ; ’
Inexact locality: Namaqualand distr., between Kamieskroon and Leliefontein,
ca. 820 m, Salter 1500 (BOL, K, PRE).
COLLECTIONS
““Galium spurium-aparine complex’’ a mean,
CAPE—3119 (Calvinia): Soetwater, 21 m W of Calvinia (-AD), Maguire |
(NBG).
276 Journal of South African Botany
—— 3129 (Port St Johns): Port St Johns, ca. 5 m (-DA), Schonland 4064 (PRE).
———3319 (Worcester): Worcester, Karoo Garden (-CB), Barker 5916 (NBG);—,
Veld reserve, Van Breda 55 (PRE), Olivier 152(a) (PRE).
3320 (Montague): Klein Roggeveld, 1 100 m (-BA?), Marloth 9600 (PRE);
Bonnievale, ca. 500 m (-CC), Levyns 4585 (BOL).
——3321 (Ladismith): Seven Weeks Poort, ca. 550 m (-AD), Levyns 2507 (BOL);
Hassaquaskloof (= ? Attaquaskloof) (-DD), Zeyher 2722 (S, SAM).
3322 (Oudtshoorn): Wilge R., WSW of Uniondale, ca. 600 m (-DB), Acocks
20619 (PRE).
EE 335 (Port Elizabeth): Hoeksplaas, ca. 1 850 m (-AB), Acocks 23532 (PRE).
3419 (Caledon): on the banks of the Zonder-Einde (Riviersonderend) (-BA),
Zeyher 2723 (SAM).
—3420 (Bredasdorp): Farm Napky, ca. 120-150 m (-BA), Taylor 3777 (PRE),
Acocks 22593 (K, PRE).
Uncertain or inexact localities: on the banks of the Olifants R. and at
Brakfontein, Ecklon & Zeyher s.n. sub PRE 21018 (PRE), Drege ‘‘76.9’’ (E);
mtns. nr. Brakfontein, Ecklon & Zeyher 2329 (SAM); on the banks of the
Kenkorivier (?spelling), Zeyher 2724 (K, S, SAM); between Stormsvlei and
Bonnievale, Acocks 1746 (S).
ACKNOWLEDGEMENTS
The present study would have been much less complete, had I not had
invaluable assistance from numerous professional and amateur botanists while in
South Africa between March 1976 and February 1977. Of all those who offered
help in various ways, I would particularly like to thank Dr O. M. Hilliard
(Pietermaritzburg) for her continued manifold support while at the University of
Natal, to Dr J. P. Rourke (Kirstenbosch) for advice and help while at the Cape, to
Mr B. L. Burtt (Edinburgh) for many useful suggestions, and last but not least to
Prof. Dr Ehrendorfer (Vienna) for constructive criticism and providing me with
unpublished data.
For assistance at the SEM and processing of the photographs I am greatly
indebted to Mrs M. G. Gilliland and Mr V. J. Bandu (EM-Unit, University of
Natal). Finally, I would also like to thank the directors and curators of all herbaria
whose material has been investigated on the spot or borrowed for their kind
cooperation.
I am greatly indebted to Drs Hilliard and Rourke for critically reviewing the
manuscript.
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Acocks, J. P. H., 1975. Veld types of South Africa. 2nd Ed. Mem. bot. Surv. S. Afr. 40.
ADAMSON, R. S. and SALTER, T. M., 1950. Flora of the Cape Peninsula. Cape
Town: Juta & Co.
The Genus Galium L. (Rubiaceae) in Southern Africa aif
BRENAN, J. P. M. et Cott., 1953. Plants collected by the Vernay Nvasal d
expedition of 1946 (part 2). Mem. N.Y. bot. Gdn 8: 406-506, y Nyasalan
CUFODONTIS, G., 1965. Enumeratio Plantarum Aethiopiae Spermatophyta. Bull.
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DréEGE, J. F., 1843. Zwei pflanzengeographische Documente. Flora 2, Beigabe.
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melten Pflanzen. Linnaea 19: 583-598.
, 1847b. Vergleichungen der von Ecklon und Zeyher und von Drege gesam-
melten sudafrikanischen Pflanzen. Linnaea 19: 599-680.
———,, 1848. Standorter-Verzeichnis, Nachtrag. Linnaea 20: 258.
Eckton, C. F. and ZeyHer, C., 1836. Enumeratio plantarum africae australis
extratropicae, pars 2. Hamburg: Perthes & Besser.
Epwarps, D. and LeIstNER, O. A., 1971.-A degree reference system for citing
pivleeical records in Southern Africa. Mitt. bot. St Samml., Miinch. 10: 501—
EHRENDORFER, F. and VERDCOURT, B., 1973. Two new species of Galium
(Rubiaceae-Rubieae) from tropical East Africa. Kew Bull. 28: 485-489.
FAGERLIND, F., 1937. Embryologische, zytologische und bestaubungsexperi-
mentelle Studien in der Familie Rubiaceae nebst Bemerkungen iiber einige
Polyploiditatsprobleme. Acta Horti Bergiani 11: 195-470.
GopLey, E. J., 1964. Breeding systems in New Zealand plants. 3. Sex ratios in
some natural populations. NZ. Jnl. Bot. 2: 205-212.
————., 1976. Sex ratio in Clematis gentianoides DC. NZ. Jnl. Bot. 14: 299-306.
Gray, S. F., 1821. A natural arrangement of British plants. Vol. 2. London.
Gigss, W., 1971. A preliminary vegetation map of South West Africa. Dinteria 4:
-114.
HOLMGREN, P. K. and KEUKEN, W., 1974. Index herbariorum. Part 1. The
herbaria of the world. Ed. 6. Utrecht: Oosthoek, Scheltema & Holkema.
KIRCHNER, O., 1911. Blumen und Insekten. Leipzig-Berlin: B. G. Teubner.
LEISTNER, O. A. and Morris, J. W., 1976. Southern African place names. Ann.
Cape prov. Mus. 12: 1—565.
Lioyp, D. G., 1973. Sex ratios in sexually dimorphic Umbelliferae. Heredity,
Lond. 31: 238-249.
Martotu, R., 1932. The Flora of South Africa 3. Sect. 2. Cape Town: Darter
Bros. & Co.
Moore, R. J., 1975. The Galium aparine complex in Canada. Can. J. Bot. 53: 877-893.
MULLER, H. 1873. Die Befruchtung der Blumen durch Insekten und die
gegenseitige Anpassung beider. Leipzig: Engelmann.
Pups, E. P., 1917. A contribution to the Flora of the Leribe plateau and
environs. Ann. S. Afr. Mus. 16: 1-379.
Purr, C., 1974. Biosystematik der Formenkreise um Galium palustre L. we
trifidum L. (Rubiaceae) auf der Nord- und Stidhemisphare. Diss. phil. Fak.
Univ. Wien.
——_, 1975. Morphologie, Biologie und Abgrenzung von Galium L. sect. Apa-
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, 1976b. Uber die Galium palustre-Gruppe (Galium sect. Aparinoides, Se
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278 Journal of South African Botany
, 1977. Chromosome numbers of some Southern African Rubiaceae-Rubieae.
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SALTER, T. M., 1937. Notes on some species in the family Rubiaceae in the Cape
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SCHREBER, C. D., 1771. Spicilegium florae Lipsicae. Leipzig: Dyk.
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Ver. vaterl. Naturk. Wiirtt. 88.
The Genus Galium L. (Rubiaceae) in Southern Africa 279
TAXA TO BE EXCLUDED
Galium horridum Thunb., Fl. Cap. 1: 556 (1813).
= Rubia horrida (Thunb.) Puff in Kew Bull. 32: 432 (1978).
INDEX OF TAXA
Page Page
Galium amatymbicum Eckl. & Galium monticolum Sond. ....... 243
Gy Dey cia an ee eases ee ECs 245 mucronatum Thunb. ........ 228, 230
aparine sensu Sond., nonL. ... 271 var. densiflorum Eckl. &
Gyp@Atie WNW, ooaccocgsbcd. 263 TAS AUNIUA cliente) aero eRe 22923
asperum Thunb. var. villosum var. subglabrum Eckl. &
IDG dk (2 AN oe cceoeeueaes 263 LEXI a6.6.06000900000 a0 yy ANS
PAGLAGHT IDE \WAGE soseooqose 255 mucroniferum Sond. .......... 259
brajjacwHicine epee oeero oe PN, BLP var. dregeanum (Sond.) Puff . 260
bredasdorpense Puff .......... 244 var. mucroniferum.......... 259
bussei K. Schum. & K. Krause . 246 namaquense Eckl. & Zeyh. . .229, 234
We? DURGGIs cesiee on aoe oot 247 namaquense Schlechter........ 263
var. glabrum Brenan ........ 247 natalensesROUY esse eee. ene Dy, pep)
CESETRS Ny Goaccocecduguc 228 rotundifolium L. var. hirsutum
SSP CAPCHSC) sis saraie se fersis ictorencres 230 Sond: Fie aconnoaceeoes YN 222
ssp. garipense (Sond.) Puff... 236 rotundifolium L. var. normale 221, 222
WEI GAINDSISS oo ccece0s0c 236 rotundifolium sensu Sond., non
var. wittbergense (Sond.) Di i Re Ce eee Appl
JEAVI TEs Gace BRO Se Rcmne 237 rotundifolium sensu Thunb.,
ssp. namaquense (Eckl. & FiO) 018) Crain aaa ehcp er chp. pie a ioO%0 221, 222
Hey bs) iRise eecrce cc 234 scabrelloides|Ruffesa-e eee 250
capense Thunb. var. expansum scabrellum K. Schum.......... 225
(@inunb)) Sond tee eases <1. 228, 230 spurium L. ssp. africanum Verde. 271
capense Thunb. var. minus stenophyllum auct., non Bak. .246, 247
Oni Sie: Metacers ais siete Siocaiw layers 2293 stenophyllum Bak. var. flavo-
capense Thunb. var. scabrum viride Utzschn. & Merxm. .246, 247
Sond eres acs oo 229. 23.4 SubyillosumiSond ear cic 255
chloroionanthum K. Schum..... 269 var. subglabrum Puff........ 257
dayscarpum Schweinf. ......221, 222 WER, SUNOS socsoncsscs 257
drescanunl Sond:) = 22-54" 259, 260 thunbergianum Eckl. & Zeyh... 221
expansum Thunb. ..........228, 230 var. hirsutum (Sond.) Verde. . 222
var. elongatum Eckl. & var. thunbergianum ......... 222
ZED se OC Re ETE 229,230 tomentosum Thunb..........-- 3
gartipense Sond). = --.-----..-229) 236 uncinatum S. 18, GIy oconeddcn Zoe
aulifiie With) tossceeaeoeac 263 uncinatum Lichtenst.........-- Ber
glabrum sensu Sond., non UndulatumeeUtieneeeeere ret: 262
hun ee es 262 wittbergense Sond. .......-- 229, zy
horridum sensu Eckl. & Zeyh., Var. glabrum Phillips Boaecnen ie
AOuPMNoy o.ccoso eo omees 271 Rubia lanata Dinter.........----
JIS. Afr. Bot. 44 (3): 281—289 (1978)
PHYTOSOCIOLOGICAL STUDIES ON TABLE MOUNTAIN, SOUTH
AFRICA:
1. THE BACK TABLE
J. GLypuHis, E. J. MOLL and B. M. CAMPBELL
(Department of Botany, University of Cape Town)
ABSTRACT
The Braun-Blanquet phytosociological technique was used to describe the vegetation of
a portion of the Back Table of Table Mountain. The fynbos vegetation was found to consist
of two communities, one containing five sub-communities, while some patches of wet
Kloof Forest occurred in well protected sites. Environmental factors, notably the moisture
régime, were found to be important in determining the distribution of the plant communi-
ties.
UITTREKSEL
FITOSOSIOLOGIESE STUDIES OP TAFELBERG, SUID-AFRIKA:
1. DIE AGTERTAFEL
Die Braun-Blanquet fitososiologiese tegniek was gebruik om die plantegroei van ’n
gedeelte van die Agtertafel van Tafelberg te beskryf. Dit het geblyk dat die fynbos
plantegroei uit twee gemeenskappe bestaan, die een met vyf subgemeenskappe. Lappe klam
kloofbos is op goed beskutte plekke gevind. Omgewingsfaktore, hoofsaaklik die beskik-
baarheid van water, is as belangrike bepalende faktore vir die verspreiding van plantegroei
gevind.
INTRODUCTION
The Back Table of Table Mountain (see Fig. 1) is covered by relatively
unspoilt fynbos vegetation, when compared to the rest of the Mountain or even the
entire Cape Peninsula.
The Back Table plateau is about 600 to 700 m above sea-level and about 200
ha in area (Fig. 2). It is divided by Disa Gorge and two reservoirs (Hely-
Hutchinson and Woodhead).
It was decided that a phytosociological survey would contribute to an under-
standing of the vegetation of the Back Table, and that this survey would also be of
use in a concurrent project concerned with the management and ecological status
of the vegetation of the whole of Table Mountain (Moll and Campbell, 1976).
CLIMATE ”
Rainfall records from Table Mountain indicate that the highest annual precipi-
tation on the Mountain occurs at Table Mountain House on the Back Table
Accepted for publication 3rd November, 1977.
281
282 Journal of South African Botany
BiGawle
Oblique aerial view, facing North, showing the Western escarpment on the left and Disa
Gorge dividing the Back Table on the right-hand side. The two reservoirs and dense Pinus
pinaster stands are just visible above the gorge.
(Schulze, 1965). The 1 780 mm annual precipitation comprises mainly winter
rainfall, though mist in summer is frequent. The western slopes and Camps Bay by
comparison receive 618 mm per annum, and Kirstenbosch on the eastern side
receives 1 413 mm (see Fig. 3).
The mediterranean climatic zone of the Southwestern Cape with cool, wet
winters and characteristically hot, dry summers, means that the greatest water
shortage and heat stress on plants is imposed during the months January to March.
GEOLOGY AND SOILS
The underlying rock on almost the entire Mountain is Table Mountain
Sandstone (Du Toit, 1954). Bedrock is often exposed. Soils vary in colour and are
generally of a sandy consistency with little humus, though occasionally soil litter
is locally abundant.
METHODS
The survey was carried out during the months March to June 1975 and for
purposes of this survey only perennially identifiable species were recorded.
Forty-eight relevés were sampled using the Braun-Blanquet technique as de-
scribed by Werger (1974). Optimal plot size was considered to be 5 X 5 m for
vegetation of a metre or less in height. Occasionally larger plots, 5 x 10 m and 10
Studies on Table Mountain: 1. The Back Table 283
x 10 m (e.g. in Kloof Forest), were used. All data collected in the field (both
environmental and floristic) were entered on a pre-printed field data sheet.
KEY
< RESERVIOR Poe /gMATABLE MOUNTAIN
s—s CABLEWAY
TWELVE APOSTLES *
YI
RONT TABLE/
ey
WESTERN TABLE | &
EASTERN TABLE: .
KIRSTENBOSCH ~
LIONS HEAD . ~~
NEWLANDS
ORANGE KLOOF
RHODES ESTATE *-
DEVILS PEAK *
CECELIA
BOUNDARY OF
NATIONAL MONUMENT
RESIDENTIAL OR aa
BUSINESS AREA
aaee2.-= ca
IDS RR SSSEEeeeeess,Y aeee
eS
Fic. 2. ,
Topographic map of Table Mountain National Monument showing the location of the Back
Table. Inset shows the position of Table Mountain on the Cape Peninsula.
The floristic data were sorted using a group-average method based on the
Canberra similarity coefficient (Campbell and Moll, 1976). This was used to obtain
televé groups. The species were then subjectively re-arranged, and a oe,
phytosociological table was constructed (Table 1). This classification was checke
in the field and found to be consistent.
284
KEY
@ cuimatic station
0” ISOHYETS.
KIRSTENBOSCH
200
Journal of South African Botany
40
CITY
HOSPITAL
00
600
100
00
CONSTANTI
FIG. 3.
LEGEND
CONTOURS — 800 —
STREAMS
BOUNDARY OF
NATIONAL MONU.
RESIDENTIAL OR
aaggaea BUSINESS AREA
———
MENT
are
KM
Map of the Table Mountain National Monument showing the major rainfall isohyets. Inset
climate diagrams indicate the duration and intensity of wet and dry seasons for two stations
representative of the climatic extremes of the area.
CLASSIFICATION OF THE VEGETATION
Three major plant communities occur on the Back Table:
1. Ilex-Blechnum Forest Community
This contains all the typical members of wet kloof forests in the Southwestern
Cape, i.e. Cunonia capensis, Ilex mitis and Halleria lucida as canopy trees, and
Maytenus acuminata, Podocarpus latifolius and Olea capensis as sub-canopy
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Studies on Table Mountain: 1. The Back Table 285
trees. The ground layer has a high cover of Blechnum attenuatum in relevé 55
(situated in Disa Gorge), and Schoenoxiphium lanceum in relevé 48 (situated in a
deep rock crevice east of Disa Gorge). Associated with both these relevés is
Asparagus scandens which is generally ubiquitous in forest (Campbell and Moll,
1977). There is a constant stream of water through the gorge draining from
Woodhead Reservoir, hence the presence of B. attenuatum and Alsophila capensis.
It is disturbing to note that Acacia melanoxylon, a pestplant (Moll and Campbell,
1976), was also recorded here.
2. Penaea-Erica Fynbos Community
The five sub-communities of the Penaea-Erica Fynbos Community all contain
one or more of the following five species: Penaea mucronata, Erica hispidula,
Watsonia spp., Hypodiscus aristatus and Clutia polygonoides. This community
occurs on both shallow and deep soils with little or much rock cover. With the
exception of sub-community E, which is essentially a wetter sub-community, the
Penaea-Erica Fynbos Community occurs on those seasonally moist to wet soils
that show some flow of water through them. It is the most extensive community
on the Back Table and varies from having a high component of forest species
present where there is some protection from fire, to having a dry component
present on well drained, exposed sites.
(i) Sub-Community A has as differential species such forest elements (Camp-
bell and Moll, 1977) as Rhus lucida/R. mucronata and Myrsine africana, riverine
Fic. 4.
Sub-community C showing Tetraria thermalis and Watsonia spp. as well as tussocked
restios such as Hypodiscus aristatus.
286 Journal of South African Botany
elements such as Myrica humilus/M. diversifolia, and Psoralea aphylla, and
fynbos elements such as Leucadendron strobilinum and Protea cynaroides.
Because of the presence of the forest elements the sub-community is placed
adjacent to the [/ex-Blechnum Community on Table 1.
(ii) Sub-community B is a depauperate form of sub-community A, probably as a
result of more frequent burning as indicated by the presence of such species as
Tetraria thermalis and Bobartia gladiata (Fig. 4). Both sub-community A and B
occur on shallow to deep, rocky soils.
(iii) Sub-community C is apparently intermediate between sub-communities A
and B, the wetter sub-community E, and sub-community D which is the driest
sub-community of the Penaea-Erica Fynbos Community (Fig. 7).
Bedrock is exposed in all relevés except 180, which had a high cover
abundance of Pinus pinaster, a non-indigenous tree species. Relevé 180 had only
14 species (average for C is 22,5) and it is suggested that relevé 180 may have
been in sub-community A if P. pinaster was absent.
The species that distinguish sub-community C as a nodum in the phytosociolo-
gical table are essentially those that dominate with frequent burning, namely
Tetraria thermalis, Bobartia gladiata, Corymbium glabrum and Psoralea acu-
leata.
(iv) Sub-community D is a drier form of sub-community C occurring on
shallower, rocky, well drained soils in exposed situations. The differential species
cee ee Fic. 5.
Cliffortia ruscifolia and Lampranthus falciformis on shallow, rocky, well-drained soils.
Studies on Table Mountain: 1. The Back Table 287
is Lampranthus falciformis, and Cliffortia ruscifolia is also present in every
televé (see Fig. 5).
(v) Sub-community E occurs on wetter sites with relatively deep soils. Two
variations are apparent, one on soils with a low rock cover, and one on soils with
an intermediate rock cover. This sub-community is related to the second fynbos
community recognised on the Back Table, namely the Erica mollis Fynbos
Community, but because of the presence of such species as Penaea mucronata,
Erica hispidula and Hypodiscus aristatus it is classified as part of the Penaea-
Erica Fynbos Community.
3. Erica mollis Fynbos Community
The differential species of this community are Erica mollis, Restio compressus
and Bulbinella floribunda. The community occurs on deep, wet, poorly drained
soils. Where the drainage is better this community abutts on to the wetter variation
of sub-community E (i.e. E”) of the Penaea-Erica Community and species which
are common to the two communities are Hypolaena crinalis (Fig. 6) and Elegia
thyrsiflora. Some species, such as Villarsia ovata and Chondropetalum
mucronatum, are even more widespread and also occur in the drier variation of
sub-community E (i.e. E’).
Fic. 6. ;
Hypolaena crinalis with Chondropetalum mucronatum (large tussocks) in a wet area.
the pines in the background.
Note
288 Journal of South African Botany
CONCLUSIONS
1. The Braun-Blanquet phytosociological technique and the computerised
sorting method of Campbell and Moll (1976) are well suited to the survey of
fynbos vegetation on the Back Table.
2. The apparently uniform fynbos vegetation of the Back Table was found to
be composed of two communities; one containing five sub-communities.
3. These communities and sub-communities are linked to environmental
factors, especially the moisture régime, as may be expected on a plateau where
aspect is not important.
4. The relationships between the major plant communities and the environmen-
tal factors are best summarised diagrammatically (Fig. 7).
DRY (well drained soils)
SHALLOW, ROCKY SOILS
pe ee S——
- ~
- ~
a z A SS
’
x — 4
Sa BT Pins ice AG
GP SeSeS
N
Z \
4 \
Vi ? Sub-community c N |
=
a,
~
\
\
\
\
\
\
\
\
4
ay
/,
/
{2
ANN
\\
\
! |
|
|
|
1
/
/
/
/
U,
a
’ Sub-community af ee )
l Le Se, ees Z i
see ene FSS eee ES
ie Se ‘i \
ul }
i 1 \ Sub-communi
N \ Sub-community E? / \ ant tyee WY
SOs ‘ \ x
LS — a
~ 5s NS
VA See bet = S Penaea-Ertea Fynbos Community ae =- aN
/ 7
Ertea mollis [
Fynbos Community Ilex-Bleehnum
Kloof Forest Community
/ \ / §
Se |
WET (free draining)
SHALLOW OR DEEP SOILS
WET (impeded drainage)
DEEP SOILS
FIRE PROTECTED
Fic. 7. .
Ordination showing the relationships between the three communities and five sub-
communities, and the major environmental factors on the Back Table.
Studies on Table Mountain: 1. The Back Table 289
ACKNOWLEDGEMENT
The authors would like to thank Miss E. Esterhuysen of the Bolus Herbarium
for many identifications.
REFERENCES
CAMPBELL, B. and MOLL, E. J., 1976. On numerical methods for classifying relevés
collected in Braun-Blanquet phytosociological surveys. JI S. Afr. Bot. 42 (1):
45-56.
CAMPBELL, B. and Mo iL, E. J., 1977. The forest communities of Table Moun-
tain, South Africa. Vegetatio 34 (2): 105-115.
Du Toit, A. L., 1954. The Geology of South Africa. London and Edinburgh:
Oliver and Boyd.
Moti, E. J. and CampBELL, B. M., 1976. The ecological status of Table
Mountain—a report on the present conservation status and recommendations for
the future management of the National Monument. Cape Town: Dept of Botany,
University of Cape Town. ISBN 0 7992 0174.
SCHULZE, B. R., 1965. Climate of South Africa. Part 8: General Survey. Pretoria:
Government Printer. 330 pp.
Wercer, M. J. A., 1974. On concepts and techniques applied in the Zurich-
Montpellier method of vegetation survey. Bothalia 11 (3): 309-323.
Jl S. Afr. Bot. 44 (3): 291-295 (1978)
PHYTOSOCIOLOGICAL STUDIES ON TABLE MOUNTAIN, SOUTH
AFRICA:
2. THE FRONT TABLE
D. LAIDLER, E. J. Mott, B. M. CAMPBELL and J. GLYPHIS
(Department of Botany, University of Cape Town)
ABSTRACT
The Braun-Blanquet phytosociological technique was used to describe the vegetation
of the study area. Two communities, one with two sub-communities and an ecotonal
community between the two sub-communities, are recognised. The siting of the Cableway
station, with the attendant large volume of tourist traffic, has led to serious vegetation
degradation in the immediate vicinity. Some recommendations as to how this situation may
be improved are given.
UITTREKSEL
FITOSOSIOLOGIESE STUDIES OP TAFELBERG, SUID-AFRIKA:
2. DIE VOORTAFEL
Die Braun-Blanquet fitososiologiese tegniek was gebruik om die plantegroei te beskryf.
Drie gemeenskappe, gebaseer op omgewingsfaktore en flora word erken. Die plasing van
die Sweefspoorstasie met die gepaardgaande groot volume besoekersverkeer het tot ernstige
agteruitgang van die plantegroei van die area gelei. Voorstelle word gemaak om die
toestand te herstel.
INTRODUCTION
The study area was the Front Table of Table Mountain, above the 1 000 m
contour (see Fig. 2. in Glyphis, Moll and Campbell, 1978). The plateau is roughly
triangular in shape with the long axis orientated east-west, about 3 km long. A
geological fault, Platteklip Gorge, divides the plateau into an Eastern and a
Western Table.
GEOLOGY AND TOPOGRAPHY
The underlying rock of the plateau is Table Mountain Sandstone (Du Toit,
1954). Between exposed areas of bedrock and boulder formations a coarse white
or grey sand overlies a dark humic soil; combined these two layers average
approximately 200 mm in depth. ;
Drainage is variable with the flat areas near the plateau centre becoming
water-logged during the rainy season, while those on the periphery of the plateau
Accepted for publication 3rd November, 1977.
291
292 Journal of South African Botany
poorly drained rock beds with dark grey, or black soils to depths of about one
metre; these are usually wet, or at least moist, throughout the year.
CLIMATE
Table Mountain lies in a mediterranean climatic region, characterised by hot,
dry summers and cool, wet winters. The Front Table has a mean annual rainfall of
1 440 mm, recorded at the Upper Cableway station (Schulze, 1965). Mists, carried
over the plateau by the summer south-easterly winds, were shown by Marloth
(1903, 1905) and Nagel (1956, 1962) to be a very important additional source of
moisture. The winter rainfall and summer mists, therefore, ensure that the plateau
has almost year round precipitation (see Fig. 3 in Glyphis, Moll and Campbell,
1978).
Temperature variation over the whole Mountain, in different seasons, is not
pronounced. On the plateaus (front and back) the mean monthly temperature
varies from 18 °C in summer to 7 °C in winter (Schulze, 1965).
METHODS
The Braun-Blanquet phytosociological technique, as described by Werger
(1974), was used in this survey. Site selection and distribution of relevés was
subjective. Only perenially identifiable species were recorded. A relevé area of
5 x 5 m was used and was found to be adequate as few species were found
outside each plot. A total of thirty-eight relevés were sampled and the floristic
and environmental data collected were entered on a standard field-sheet for each
relevé.
Using a computer programme coefficients of similarity for each relevé, based
on floristic data, were calculated (Campbell and Moll, 1976). From this informa-
tion a dendrogram, showing the floristic relationship of the relevés, was drawn.
Dendrogram groups, with certain minor adjustments, were used to construct the
final phytosociological table. Species sequence in the table was arranged subjec-
tively (Table 1).
DISCUSSION
Description of the Communities
1. Maytenus—Athanasia Community
The single relevé (140) is provisionally used to describe this community,
which occurred on a rocky, well-drained, steep cliff where the soil was dark grey
to black, and from 100 to 300 mm deep. The relevé itself was sited near the
Upper Cableway station and contained, in addition to the dominants Maytenus
oleoides and Athanasia parviflora the following species, Salvia africana, Myrsine
africana, Peucedanum galbanum, Cliffortia ruscifolia, Anthospermum aethiopi-
cum and Ehrharta ramosa.
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Studies on Table Mountain: 2. The Front Table 293
2. Restio Community
The major physiognomic component of the upper plateau is a closed Restio
Community less than 1 m tall, in which Chondropetalum ebracteatum, Ursinia
nudicaulis, Ehrharta setacea, Chondropetalum mucronatum, Villarsia ovata and
Restio sarcoclados are the differential species. Two major sub-communities,
separated by a fairly well-differentiated ecotonal group of species (G) occur
within the Restio Community.
(i) Watsonia-Penaea Sub-community (A to F). This sub-community occurs on
the shallow, essentially pallid soils overlying bedrock (see Table 2) and has been
burnt fairly frequently in recent years. Important differential species, apart from
Watsonia spp. and Penaea mucronata, are Cliffortia ruscifolia, Thamnochortus
nutans and Erica hispidula. It is noteworthy that T. nutans, which is common on
the upper plateau, has an extremely limited distribution in the Fynbos Biome as a
whole, being restricted to the top of Table Mountain and the top of the
Constantiaberg on the Cape Peninsula.
Within the sub-community there occur a number of variations, these are
discussed below:
The Restio-Schizaea variation (B) occurs on the temporarily moist, relatively
steep sites and has as differential species Restio perplexus, Schizaea pectinata,
Aristea spp., Cassytha ciliolata and Anemone capensis. On wetter, flatter sites,
with shallow soils, Simocheilus depressus and Erica plukeneti which also occur in
this variation, are able to grow successfully indicating an ecotonal type (A). On
somewhat deeper, pallid soils that have not been burned for five or seven years
Tetraria cuspidata and Leucadendron strobilinum are more common (C), and
where fire has burned the vegetation recently Euryops abrotanifolius/Ursina sp.
and Elegia spp. occur, but Erica coccinea and L. strobilinum are absent (D).
On those shallow, pallid soils that have a high surface rock cover a Centella
variation occurs (E). Differential species of this variation, apart from Centella
caespitosa, ate Helichrysum pinifolium, Chondropetalum deustum and Anaxeton
arborescens. On flatter sites where the surface rock cover is less Centella does not
occur and Restio bifidus and Chrysothrix capensis form an ecotonal variation (G)
between the Watsonia-Penaea Sub-community and the Restio-Hypolaena Sub-
community. Here the topography is relatively flat, soils deeper, rock cover is low,
and Restio bifidus, Chrysothrix capensis, Afrachneria capensis and Stoebe incana
are more frequent. i
(ii) Restio-Hypolaena Sub-community (H & I). This sub-community occurs on
deeper, essentially pallid, seasonally wet soils on relatively flat sites. Differential
species are Hypolaena crinalis, Restio compressus and Senecio CTESDUS, Two
variations, one characterised by Elegia thrysiflora, Erica sp., P entachistis palles-
cens and Berzelia lanuginosa on the deepest soils (I), and the other characterised
by the absence of these species and high cover abundance values of H. crinalis,
occur (H).
294 Journal of South African Botany
Anthropogenic influences
The location of the Upper Cableway station on the Western Table is respon-
sible for a large volume of tourist traffic on the Front Table. This has resulted in
extensive degradation of the vegetation especially within close proximity of the
station and restaurant. Further away there is also much damage and despoilment
along the edges of the foot-paths particularly along the front face.
It is, therefore, strongly recommended that well landscaped paths be con-
structed in areas where degradation is visible. This would, in fact, entail the
planning of the path network on the whole Western Table, so that a few major
routes are hardened. Visitor access should then be restricted to these paths to
prevent further extensive degradation of the plant communities and allow their
recovery.
CONCLUSIONS
Two distinct plant communities are recognised and described using the
Braun-Blanquet phytosociological technique. One community, the Restio Com-
munity, has two sub-communities and an ecotonal variation. The important
environmental factors determining these communities are soil depth, moisture
content and rock cover. The trampling of the area by visitors to Table Mountain is
causing concern regarding the future conservation status of the Restio Com-
munity.
ACKNOWLEDGEMENT
The authors would like to thank Miss E. Esterhuysen of the Bolus Herbarium
for many identifications.
REFERENCES
CAMPBELL, B. M. and MOLL, E. J., 1976. On numerical methods for classifying releveés
soled in Braun-Blanquet phytosociological surveys. JI S. Afr. Bot. 42(1):
5-56.
Du Toit, A. L., 1954. The Geology of South Africa. London and Edinburgh:
Oliver and Boyd.
Grypuis, J., Mort, E. J. and CampBeLt, B. M., 1978. Phytosociological studies
on Table Mountain, South Africa: 1. The back Table. JUS. Afr. Bot 44 (3): 281-289.
MarLotH, R., 1903. Results of experiments on Table Mountain for ascertaining
the amount of moisture deposited from south-east clouds. Trans. S. Afr. phil. Soc.
14: 403-408.
MARLOTH, R., 1905. Results of further experiments on Table Mountain for
ascertaining the amount of moisture deposited from the south-east clouds. Trans.
S. Afr. phil. Soc. 16: 97-105.
NAGEL, J. F., 1956. Fog precipitation on Table Mountain. Q. Jl. R. met. Soc.
82(354): 452-461.
~~ 4
Studies on Table Mountain: 2. The Front Table 295
NaGEL, J. F., 1961. Rainfall in the Table Mountain area with special reference to
orographic effects. Weather Bureau Notos, Pretoria 10: 59-73.
ScHULZE, B. R., 1965. Climate of South Africa. Part 8: General Survey. Pretoria:
Government Printer. 330 pp.
WerGER, M. J. A., 1974. On concepts and techniques applied in the Zurich-
Montpellier method of vegetation survey. Bothalia 11(3): 309-323.
Ji S. Afr. Bot. 44 (3): 297-312 (1978)
AN INVESTIGATION INTO THE ECOLOGY AND PRODUCTIVITY OF
EPIPHYTIC MOSSES
N. H. G. JACOBSEN
(Transvaal Division of Nature Conservation, Pretoria)
ABSTRACT
Experimental harvesting of moss was undertaken for five years at two Transvaal
localities. Determination of productivity was the main aim. It was established that mosses
are very slow growing and require a minimum of seven to 10 years after harvesting to
regenerate. Annual production amounted to 4 700 g/ha which is only 7,64% regrowth,
while biennial plots exhibited 15,45 % regrowth and triannual plots produced 27,38 % of the
initial harvest. As Transvaal only has approximately 15 959 hectares of harvestable moss
(5 426 002 kg), which on a sustained yield basis can be divided into 2 279,83 ha yielding
775 143 kg moss/annum on a seven year basis while over 10 years, ! 595,88 ha could be
harvested with a sustained yield of 542 600 kg/annum.
UITTREKSEL
*"N ONDERSOEK NA DIE EKOLOGIE EN PRODUKTIWITEIT VAN EPIFITIESE
MOSSE
Eksperimentele versameling van mos is vir vyf jaar by twee Transvaalse lokaliteite
gedoen. ’n Bepaling van die produktiwiteit was die hoofdoel. Daar is vasgestel dat mosse
baie stadig groei en ’n minimum van tussen 7 en 10 jaar na dit geoes is, word benodig om
te herstel. Die jaarlikse produksie was 4 700 g/ha wat slegs ’n 7,64 % herstel verteenwoor-
dig terwyl persele wat elke tweede jaar geoes is ’n herstel van 15,45 % en persele wat na
drie jaar geoes is 27,38 % van die oorspronklike oes gelewer het. Aangesien Transvaal net
sowat 15 959 ha oesbare mos het (5 426 002 kg), kan dit teen ’n volgehoue basis in
2 279,83 ha gedeel word om 775 143 kg mos/jaar te lewer op ’n sewejaar basis terwyl daar
oor 10 jaar van 1 595,88 ha ’n volgehoue produksie van 542 600 kg/jaar geoes kan word.
INTRODUCTION
On account of their water holding capacity epiphytic mosses are widely
utilised by florists to pack flowers for shipment, for floral arrangements and for
exhibition purposes. This has resulted in a demand for moss, that is harvested by
Bantu gatherers and sold to the florists. The moss is harvested in the indigenous
forests of the Transvaal, mainly in the east, along the Transvaal Drakensberg but
also in the Soutpansberg. As these areas are the main source of moss in the
Transvaal and because of the limited area covered by indigenous forest, the supply
of moss is likewise limited. In addition, as large areas of forest lie in Bantu
homelands, the protection afforded them by the Department of Forestry falls away
and these areas are being severely cut back by indiscriminate woodcutting,
burning and the foraging of livestock.
Accepted for publication 15th December, 1977.
297
298 Journal of South African Botany
In view of these limitations in the supply of moss, the harvesting on a large
scale was considered with concern by the Division of Nature Conservation and
mosses were placed on the list of Transvaal Protected Plants. The demand of the
florists was however not ignored and an investigation into the ecology and
production of mosses was initiated by Mr J. Jooste of the Division of Nature
Conservation to determine the effects of harvesting, so that effective management,
with the object of a sustained yield could be instituted. Adequate permits could
then be issued to avoid over-exploitation. The aims of the project were therefore
essentially threefold:
(a) Determination of the amount of epiphytic mosses in the indigenous
montane forests of the Transvaal.
(b) Determination of the time it takes for the moss to regenerate fully after
harvesting.
(c) Determine the areal percentage which can be annually harvested on a
sustained yield basis.
METHODS
For a determination of the total amount of harvestable moss it was necessary
firstly to establish the total area in which moss occurs. Areas of indigenous forest
were calculated from existing maps and aerial photographs with the aid of a
planimeter, after local inspection to check on the distribution of moss.
In two areas, namely Woodbush and the farm Bloemfontein in the Sout-
pansberg, plots 100 square metres in extent were laid out. In the former a series of
five plots was set out initially in June 1970, of which one (E) could not be
re-located a month later (Jooste, 1971). This was followed by another set of 10
plots during July, 1972. The former set was used for annual growth checks, while
five of the latter were used for biennial checks and the remaining five for triannual
checks on regeneration. At Bloemfontein a set of five plots was set out and
harvested annually during January.
The plots were laid out in various sections of the forests so as to cover a
variety of microclimates ranging from dry to wet. The selection of plots was
arbitrary and their suitability visually estimated. Each plot was beaconed off with
numbered wood or iron poles at the corners.
During 1972 all the plots at Woodbush were subdivided into quarters,
(subplots), in order to establish whether any variations existed within each plot as
a result of climatic or other local factors.
To determine the surface area, the trees within the plots were measured to an
arbitrary height of two metres, based on the assumption that the Bantu gatherers
did not harvest beyond that point. The circumference of the trees was measured at
one metre from base and multiplied by the arbitrary height thus giving the area of
the tree trunk. In cases where the trees had large buttress roots measurements were
taken from where the trunk became vertical.
Ecology and Productivity of Epiphytic Mosses 299
The moss was harvested by hand, placed in plastic bags and then sorted to
remove foreign material such as pieces of bark and soil. It was then transferred
into paper bags, oven dried over a period of 24 hours at 85 °C (185 °F) and
weighed.
The average weight of moss per plot (100 m2?) was used to calculate the
amount of harvestable moss available.
For a determination of the time taken for moss to regenerate completely the
plots were visited and harvested annually. The two and three year plots at
Woodbush were harvested during 1974 and 1975 respectively. It must be stressed
here that harvesting during experimentation was more severe than that of the
Bantu gatherers who removed only the moss which was readily available.
RESULTS
Area determination
In the area calculations allowances were made for roads and open areas. These
calculated areas are set out in Table 1. Excluded are the indigenous forests on the
farm Bloemfontein in the Soutpansberg, in the Blyde River Canyon and in the
Sabi area. The tabulation, however, contains the major areas of forest which were
investigated.
TABLE 1.
Surface area of indigenous forest and surface covered by moss in the various plantations.
Total area of
indigenous Area in which
Area forest moss is found
(hectares) (hectares)
ELAS Lips sopra Bie = ees ee RE EE ites 206,78 190,20
VWigaali nese, Se Soe oes ote Oe nO mn 4 271,50 3 630,76
DZ ECA er Oo eS irene Seta een etic 2 355,43 2 143,46
WEaKICDSKOP Mane ne athe aie cons see ois 3 039,35 2 811,39
ETtEADE Tite eset I EI Se. SEU: oS e oe 1 711,38 1 oo 0
SEITE OSE Ss See eee ce Se a Or rece acnaD 2 819,04 2 678,0
New [CECE ee « SE RSIS EEO Re POE 2 197,24 2 087,37
IReodewal/Goedehcopes: 2.75.50 - ee ea: B38 7222 320,35
INV a SHA Be Pet stay tty ireici sree ne sai a8s iokyort 463,45 444,91
INieshontstontemileS opera seen eee 43,63 43,63
Ee ee. PE oe 15 958,83
In the Bantu homeland area of the Soutpansberg only the farm Bloemfontein
was found to have moss in harvestable quantities. The area of indigenous forest
does not cover more than about 300 hectares, parts of which are rapidly being
destroyed so that this area can probably be ruled out as a future source, especially
as illegal harvesting of moss is adding to a swift deprivation (P. W. de Wet
Wessels, pers. comm.)
300 Journal of South African Botany
The total area of indigenous forest in the Transvaal with the exception of small
areas scattered in the east and north east is thus 17 444 hectares, of which 91,49 %
or 15 959 hectares contain epiphytic mosses. This is only 0,06 % of the total land
area of the Transvaal which fortunately, however, falls mainly under the Depart-
ment of Forestry and is thus afforded some protection.
Regeneration Period
The moss is harvested manually by the Bantu gatherers to a height of two
metres. Experimental harvesting extended over the six year period 1970—1975,
though only for five years on the farm Bloemfontein, while at Woodbush the year
1971 was excluded as the initiator of the Project left the Division and the author
continued with the Project only in 1972. Annual harvesting therefore took place at
both sites for four consecutive years. Unfortunately the first year of reharvesting
resulted in some discrepancies as may be seen on Table 2. This may be due to
varying sampling techniques as it was not always possible to recognize which
trees had previously been harvested, the present author not having accompanied
the initial researcher during his visits, so that perhaps different standards were
used in determining the extent of harvesting. The greatest discrepancy lies in the
method of measuring the surface area, that is circumference and height. The
method used by the present author is noted above. When comparing the figures
from Bloemfontein it will be noticed that in plot C the difference in surface area
between 1971 and 1972 is very great and probably due to measuring error, as it is
doubtful that the growth rate on 35 trees would vary to such an extent.
When the present author took over, all the trees within the plots were
re-measured and subsequently over the following three years only those trees with
harvestable moss were measured so that moss regrowth per unit area could be
established and compared. Large differences in the weights of moss per unit
surface area such as are exhibited by plots A and C could also be due to the
harvesting of certain moss species which were not reaped by the previous worker.
As there was no request for species determination, it was thought best to harvest
all moss species on the trees within the plots and this procedure was adhered to
throughout the duration of the project. No change in species diversity was
observed throughout the duration of the study and regeneration was apparent from
the moss remnants left behind after the previous harvesting. In some instances
young plants were seen germinating in areas which were moister than the
surroundings but these seedlings appeared to be the same species as the adults
harvested on the same trunks. One of the commoner species involved was
Papillaria natalensis Sim. Florists utilise the moss irrespective of length or
species. The results of the harvesting are shown on Tables 2, 3 and 4 in the
Appendix.
From these it is evident that experimental harvesting has a considerable
influence on the regeneration of moss. Particularly noteworthy is the reduction in
Ecology and Productivity of Epiphytic Mosses 301
the number of trees with harvestable moss among the annual plots, both sites
showing similar trends in the fluctuations of average numbers of trees per plot per
annum. (Table 5(a) and (b)).
TABLE 5.
Fluctuations in average number of trees per plot per annum.
(a) Woodbush
1972 1973 1974 1975 Total Ave
Les, Cid ae ae an ae a 23 19 17 12 71 ES
PPE io estat ahets isis 30 13 22 14 79 19,75
CPP ns ees 30 9 18 15 1 18,00
1D) re-center et 17 0 9 9 35 8,75
Notale wr#. oe eee eres 100 41 66 50 257
LC SOR oe omaee 25 10,25 16,5 IDES 64,25
(b) Bloemfontein
1972 1973 1974 1975 Total Ave
|. es O REACT 25 15 21 16 Wl 19,25
Le 5 OS Oe ena 27 1 21 11 60 15,00
Cpr es. 35 0 10 8 53 13,25
1D oi Oi ee 23 1 NR NR 24 12,00
Pe aes. eer chases 17 0 0 6 23 5,75
otaleere as ee 127 7/ 52 41 PX
Vea riety. Ue nen 25,4 3,4 13,0 10,25 59,25
NR = not recorded during 1974—1975
These fluctuations are mainly due to the fact that moss regenerates very
Slowly, particularly after the first severe harvesting to which it is subjected.
Consequently when the sites are re-visited only those trees with moss of harvest-
able length, that is one centimetre or more, are measured, while those which have
moss of only very short growth are left until sufficient re-growth permits
harvesting at later visits. The fluctuations are also documented in the graphs (Fig. 1)
Showing the annual yield of moss per plot. However, an average of the
fluctuating yields will give a better picture as to the amount of regenerated moss
per year. Over the three years subsequent to 1972 the average yield of moss
harvested per plot from Woodbush and Bloemfontein amounts to 47 grams and 45
grams (dry weight) respectively (see Tables 2 and 3). In contrast the biennial plots
yielded an average of 69 grams per plot and it is noticeable that only 48,5 % of the
trees had harvestable moss after two years. The triannual plots show a substantial
increase and an average of 101 grams per plot was harvested. However, only 51%
of the trees were found with harvestable moss representing only a slight increase
on the biennial plots and the equal percentage to the average of the annual plots.
It, therefore, appears that at Woodbush approximately 50% of the trees remain
barren after being harvested and that the moss requires a longer period to
regenerate on them.
302
AMOUNT OF MOSS (kg/m?)
AMOUNT OF MOSS (kg/m2)
Journal of South African Botany
BLOEMFONTEIN
4
Jn
S,
°
°
9
8
ceage Si ems memes ee meres ee
YEAR
WOODBUSH
Fic. 1.
Annual fluctuations in the amount of moss harvested.
Ecology and Productivity of Epiphytic Mosses 303
Factors controlling Re-generation
There are a number of factors which influence the regrowth of moss. Prior to
harvesting, the moss hangs festooned from the trunks and branches of the trees as
also from rocks in the forest. Moss grows more profusely in a relatively closed
canopy forest than in an open forest where penetration by wind and sunlight
causes dehydration. The plots laid out in the forest aimed at the investigation of
these variations. This is clearly demonstrated at Woodbush when plots A and C
are compared, the former giving much larger yields than the latter. A similar
variation was observed at Bloemfontein.
Slope aspect appears to influence the growth of moss as it frequently only
grows on one side of the trunk, leaving the opposing side bare. This, no doubt, is
due to the prevailing weather conditions which bring more moisture to the exposed
side of the tree trunk while the other side remains in a partial rain shadow. Higher
up the tree the disparity disappears.
At Woodbush the E, SE and S sides of the stems have the most luxurious moss
cover. The same applies to younger trees where the bark is usually smoother than
in the adult stage and here it needs several years for the moss to establish itself.
Rough bark is more densely moss-covered because its rugosity offers innumer-
able places for the moss spores to lodge. It also collects falling dust and humus
which lodge in the cracks and form a growth medium in which the spores can
germinate, enhanced by a higher degree of water retention as compared with
smooth bark which dries fairly rapidly after rain. Harvesting moss from the former
is more difficult than from the latter as usually only the easily removable portion
of the plants is taken, whereas the moss on smooth trees frequently comes away
entirely with stolons and rootlets so that nothing is left for regeneration. Doubt-
lessly this has a great influence on the assessment of the number of trees with
harvestable moss.
Areas in which the undergrowth is relatively dense also appear to produce
more moss due to protection from wind and sunlight, while transpiration by the
shrubs and herbs also contributes to a moister microclimate. This was seen at
Woodbush where moss growing on tree trunks in the herbaceous layer was longer
than that on the same tree trunk in a more exposed position.
The influence of Moisture
There does not appear to be a well defined correlation between the amount of
moss production and rainfall, but it is felt that this is the case between the growth
of moss and the number of misty days per annum. This view is supported by the
fact that the average amount of growth of moss per plot at Woodbush and
Bloemfontein is very similar although the average rainfall at Woodbush over the
four year period 1969 to 1972 is 2,5 times as much as that at Bloemfontein.
Taking the three most favourable plots at Woodbush and Bloemfontein, a marked
similarity in the amount of regrowth of the epiphytic mosses is revealed.
304 Journal of South African Botany
Experiments undertaken by R. Marloth (Wellington, 1955) on top of Table
Mountain showed that during a period of 21 days, the rainfall gauge registered nil
whereas a gauge adapted to condense moisture from mist recorded 758,4 mm. On
a previous occasion (January-March 1904) the rain gauge also recorded nil
whereas the mist gauge registered 484,4 mm. This stresses the importance of mist
as a source of moisture especially as it occurs throughout the year in the areas
where moss grows. Condensation occuring on the leaves and boles of trees runs
down the trunks providing a direct water supply to the moss, besides a certain
amount of moisture from the mist which may be absorbed directly. This process
was noticed on the Downs in the eastern Transvaal Drakensberg where water was
seen streaming down the boles of the trees. Unfortunately there is no record of the
number of misty days per annum at either site, so that a numerical correlation is
not possible.
Both areas have a similar rainfall pattern with a pronounced peak during the
four months December, January, February and March (Table 6). Although on the
average rainfalls occur throughout the year there is a distinct minimum during the
winter months. Bloemfontein is more arid than Woodbush and usually experiences
at least one month without rainfalls. During this period (usually July) the moss
becomes dry and brittle but greens up and starts growing during August at
Woodbush. There is a great fluctuation in the amount of rain from year to year,
being in some years two to three times as much as in other years (Table 6). In
these mountainous areas it often rains for days on end with as much as three
weeks of cloudy rainy weather, while on the other hand thunderstorms of short
duration but intense rainfall occur.
Evaluation of Results
The results reveal that the initial annual harvesting yielded 0,034 kg/square
metre. This dropped radically (see Fig. 1) after the initial harvesting and the
average annual growth of moss on the plots yielded 0,002 kg/sq. metre (7,6 %). In
contrast the biennial plots yielded 0,11 kg/sq. m on the initial harvest, but two
years later only 0,017 kg/sq. metre (15,45%) had re-grown, which is just over twice
the productivity of the annual plots. The triannual plots, which when first
harvested yielded 0,029 kg/sq. metre, gave at the second harvest only 0,008 kg/sq.
metre (27,5 %). This is 3,62 times as much as that of the annual plots and 1,77
times as much as the biennial plots calculated on a percentage basis. At this
growth rate the moss would take a minimum of seven years to reach the initial
amount harvested and may even take up to 10 years.
The layout of the experimental plots lends itself to statistical analysis by means
of a randomised complete block design. The data from the annually harvested
plots (1972, 1973 and 1974) were analysed separately from those harvested
sulin, (1972 and 1974) and triannually (1972 and 1975) (see Tables 7, 8 and
).
Ecology and Productivity of Epiphytic Mosses 305
TABLE 7.
Analysis of variance of moss weights (kg per 10 x 10 m plot) collected annually in 1972,
1973 and 1974.
Source of Sum of Degrees of
variation squares freedom Mean Square F
L1G eeetendee tele aig 0,008819 | 3 0,002940 2,63
Subplots22) 22 s42 2.5 0,050897 36 0,001414 1,27
CarspeyS ie Sues cbc 0,046448 0,023224 20,81*
DANO ae ete MO Og eee 0,006693 0,001116
a Mee oe 5 0,1222857
*Significant at 1 % level.
TABLE 8.
Analysis of variance of moss weight (kg per 10 x 10 m plot) collected biennially in 1972
and 1974.
Source of Sum of Degrees of
variation squares freedom Mean Square
Blotste Same cites sa 0,029397 + 0,0007349
SUDPLOES je co -cnheser shen 0,238244 30 0,007941
iG RS aoe ae 0,197543 ] 0,197543
| EATR0t cyecsenegie ae ee anes 0,024501 4 0,006125
SLOMAN apse ¢ ycrs eyei 0,489685 39
*Significant at 1 % level.
TABLE 9.
Analysis of variance of moss weight (grams per 10 x 10 m plot) collected triannually in
1972 and 1975.
Source of Sum of Degrees of
variation squares freedom Mean Square F
RIOESRe o cree orskeic se 325,0 81,25 0,78
MEdISte cece tees 1 210,0 1 210,00 jie
EE ca 413,0 103,25
*Significant at 5 % level.
The results were on the whole similar for the annually, biennially and
triannually harvested plots. No significant variation was found to be due to the
plots or the subdivisions within each plot, indicating that conditions within each
plot and subplot were similar, such as the total surface area available for moss
growth, moisture (partly) and temperature conditions, etc. However, very signi-
ficant differences were found to be caused by the annual treatment, which were
most likely due to variations in the way of harvesting of moss from year to year.
For instance during 1972 the moss was harvested from the majority of trees within
the plots. In 1973, however, many of the trees had moss of only very short length
306 Journal of South African Botany
(5 mm or less) which could not be harvested in standard fashion and was thus left
until 1974 when it might be long enough for plucking.
Some influence is obviously due to the variation in the number of misty days
per annum at Woodbush, as it has been shown that mist contributes significantly to
the amount of moisture available to mosses.
It is interesting to note that increases or decreases in moisture and *‘treatment’’
do not seem to affect all plots to the same degree. Table 10 reflects the proportions
of moss collected from each plot in 1972, 1973, 1974 and 1975 respectively
considering plot C as unit. It will be noted that plot B produced 2,6 times as much
in 1972, 2,3 times as much in 1973, 2,9 times as much in 1974 and only 1,0 times
as much in 1975. This variation is also apparent in the other plots such as A where
the production went from 1,9 times that of C in 1972 to 5,1 times as much in 1973
and dropped to 3,3 times in 1974 while it was only 1,3 times in 1975. It could
here be noted that plot A is probably the most favourably situated plot of the set
with regard to moisture and other aspects and therefore consistently shows a
higher growth rate. Plot D situated in the most xeric situation also shows similar
trends.
TABLE 10.
Comparison of proportions of moss produced from each plot during successive years. Plot
C taken as unit in all cases.
DISCUSSION AND CONCLUSION
The average annual growth of moss at Woodbush over the three years 1973,
1974 and 1975 is 47 grams per plot while that at Bloemfontein is 45 grams
indicating that in spite of the differences in rainfall, climate and edaphic condi-
tions, similar growth is achieved. This lends support to the view that mist is an
important factor for the growth of moss. The average regrowth is 7,64 % per
annum. The biennial plots show a regrowth of 15,45 % over the two periods which
is slightly more than double that of the annually harvested plots. The triannually
harvested plots produced 27,58 % of the initial harvest after three years which is
slightly more than three and a half times as much as the annually harvested plots
and about 1,75 times as much as the biennially harvested ones. The growth rate
appears to accelerate over three years as an average rate of 8,99 % is achieved. It is
difficult at this stage to extrapolate the further rate of growth but it is assumed that
this rate will probably not be maintained. At the present rate it will take the moss a
minimum of seven years and possibly as much as 10 years to regenerate fully. It is
not known at what stage moss actually reaches maturity and when it would be
Ecology and Productivity of Epiphytic Mosses 307
most productive to harvest. It may be that the moss after reaching a certain length
slows down completely so that harvesting on a sustained yield basis may best be
done at an earlier stage when the standing crop has reached optimum growth and
will begin to slow down. This stage may already be reached at seven years after
harvesting. It must, however, be emphasised once again that harvesting during the
experiment was considerably more severe than that of the Bantu gatherers who
take only what is readily available, leaving ample moss and thus probably
speeding up the process of regeneration.
Based on the average amount of moss initially harvested from the plots, 340
kg could be harvested per hectare which when calculated to cover the 15 958,83
hectares amounts to an overall weight of 5 426 002 kg of moss. In order to
manage the reaping of moss on a sustained yield basis over a period of seven years
2 279,83 hectares could be harvested giving 775 143 kg of moss per year. On a 10
year rotational basis 1 595,88 hectares could be harvested giving a sustained yield
of 542 600 kg per year. While the present experiment only sampled the moss
growing on the trees itself, mention should be made of the large amounts also
found hanging from the branches of smaller trees, on dead logs and on rocks
which contribute considerably to the standing crop of a hectare of forest. This is,
however, not uniform and some patches of forest are devoid of this. In addition it
appears that once the moss has been removed from dead or decaying wood no
subsequent regrowth occurs apart from that left from the previous plucking. There
is, therefore, at least initially considerably more moss than the above figures
indicate. In conclusion it may be worthwhile to harvest a set of plots after a period
of five years and again after seven years, which will then give a good indication of
the rate of growth during these periods as well as determine the best time interval
between harvesting. The moss itself, however, limits the interval between harvests
and is, therefore, the main limiting factor.
ACKNOWLEDGEMENTS
The following people and institutions are thanked for their help and advice:
The Departments of Forestry, Bantu Administration and Development for their
generous consent to work in the areas under their control.
Mr. P. W. De Wet Wessels for rainfall data from the farm Piesanghoek,
Soutpansberg District, as well as for allowing the author to camp there.
Dr. S. M. Hirst for help and guidance in the continuation of this project.
REFERENCES ee
JACOBSEN, N. H. G., 1974. An investigation into the ecology and productivity of Epiphytic
mosses. (Progress report (1974)). T.P.A. Nature Conservation Division. og
Jooste, J. P., 1971. Ondersoek na die ekologie en produktiwiteit van Epifitiese
mosse. Flora Navorsing: Projek T.N. 6/4/3/4. Afdeling Natuurbewaring T.P.A.
WELLINGTON, J. H., 1955. Southern Africa: A geographical Study. Vol. 1. p. 260.
London: Cambridge University Press.
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JIS. Afr. Bot. 44 (3): 313-315 (1978)
BOOK REVIEWS
DORMANCY AND THE SURVIVAL OF PLANTS, by Trevor A. Villiers. Studies in
Biology No. 57. London, Edward Arnold, 1975. 68 pp. Price £2,80 (hard cover).
This book, written by an acknowledged expert in the field of dormancy, covers ten short
chapters, dealing with: the dormant state; climate and survival; structural modifications in
dormancy; resistance of dormant organisms; control of entry into dormancy; types of seed
dormancy; metabolism during dormancy; biochemical controls and dormancy; emergence
from dormancy; and conclusions.
As the chapter headings indicate, just about every facet of dormancy worth considering
is discussed. This is done in an easily readable style, well illustrated with graphs, tables,
line drawings and schematic representations. As a well-balanced introduction to the subject
it can hardly be improved upon (bearing in mind, of course, the space restrictions applicable
to a book in this series). Particular attention is given to the role of hormones in the
induction, maintenance and termination of dormancy, in conjunction with the environmental
“‘triggers’’ involved, but I think that the views of Vegis in this regard could have received
some more attention, even in a book at this level.
This book is clearly not intended for specialists on the subject of dormancy, but even
they can benefit from its reading. It can be highly recommended for undergraduate students
in plant physiology and horticulture, and should make very useful reading for all botanists
and plant scientists in general.
J. A. DE BRUYN
COMPOSITAE IN NATAL, by O. M. Hilliard, with pp. xi+659 incl. 25 pp. line
drawings. Pietermaritzburg: University of Natal Press, 1977. R24,00. (Orders to:
University of Natal Press, P.O. Box 375, 3200 Pietermaritzburg, South Africa.
American and Canadian orders to: Laurence Verry, Inc., Mystic, Connecticut
06355, U.S.A.).
More than a century has elapsed since the appearance of Harvey’s classical treatment of
the Compositae in Flora Capensis. Perhaps another century will have gone before the
ambitious, new Flora of Southern Africa is finished. Under these circumstances a modern
regional account of a large family is most welcome. Dr O. M. Hilliard’s Compositae in
Natal must be added to the list of standard works on the South African flora. The present
volume also paves the way for the complete treatment of the family in the new flora.
As someone who has studied the South African Compositae, particularly in Natal, for a
long time now, Dr Hilliard was certainly the right person to undertake such a comprehen-
sive task. During many years of field work she has discovered numerous undescribed
species and many new records for Natal. Moreover, she has critically revised the material in
herbaria both at home and overseas, a most laborious task. Part of the original research
which preceded this volume has been published in a series of joint papers with B. L. Burtt
(see: Notes Roy. Bot. Gard. Edinb., 1970—). '
Not surprisingly, the Compositae is the largest family of flowering plants found in Natal
province. The present account deals with no fewer than 640 species, belonging to’ 113
genera. They include constituents of most of the vegetation types present in the province,
covering a wide range of habitats from littoral scrub to the alpine heaths of the highest
mountains in South Africa. Weeds and aliens are also covered, with notes on their origins
and other useful information. The text provides determination keys, full generic and specific
descriptions, notes on geographical distribution, ecology and flowering times, and frequent
a3
314 Journal of South African Botany
taxonomic comments, e.g. on infraspecific variation and nomenclature. Although the
generic descriptions are modelled on those of Bentham, the sequence adopted is that of
Dalla Torre and Harms. The precise and accurate specific descriptions are all original. All
types seen by the author are fully cited and the relevant synonyms given, also with types.
The primary aim of the book is to aid accurate identification, for which purpose good
alternative keys are given. There is a general key to all genera, but one can also proceed via
the key to the tribes followed by the generic keys for each tribe. There are also keys down
to species level for all genera which include more than one species.
The classical 13 tribes are recognized and all are represented, although in some cases
only by introduced species (Heliantheae, Helenieae, Cardueae). The author is well aware of
the current conflict of opinions on tribal circumscription, but she has wisely adopted a
conservative taxonomic frame-work.
The line drawings are both accurate and artistically appealing, and more would have
been very welcome. A useful feature is the introductory series of drawings, explaining the
niceties of synantherological terminology. On the inside covers of the strongly cloth-bound
volume are two survey maps. A comprehensive index to all taxa, including synonyms,
completes the text. Relevant bibliographical references are given in the headings to genera
and species, sometimes also in the commentaries.
This important publication is obviously indispensable to all students of South African
Compositae, but will also be of great interest to taxonomists and phytogeographers in
general. Ecologists, agriculturists, gardeners and biology teachers will also find much useful
information in this book, which will undoubtedly attract considerabie attention far outside
South Africa itself.
BERTIL NORDENSTAM
Woop STRUCTURE IN BIOLOGICAL AND TECHNOLOGICAL RESEARCH, edited by P.
Baas, A. J. Bolton and D. M. Catling, with pp. 280, 28 fig., 25 plates. (Leiden
Botanical Series No. 3.) ISBN 90 6021 302 5. Leiden University Press, 1976.
(approx. R28,00).
Despite its title, this is a collection of studies in plant anatomy. It arose from the
proceedings of the Anglo-Dutch Wood Anatomy Meeting, organised by the Royal Micro-
scopical Society at Oxford and Kew in April 1976. Appropriately it opens with an
entertaining essay on the history of the Jodrell Laboratory, with whose centenary this
meeting coincided. The former Keeper, C. R. Metcalfe tells of the personalities and events
that led to the establishment of what might be called the English School of systematic
anatomy, at Kew. A useful survey by J. D. Brazier deals in general terms with the use of
wood anatomical features in identification and in taxonomy, both of which present their
own peculiar problems. This is well shown in the two detailed papers which follow. T.
Baretta-Kuipers discusses the taxonomic position of the genera, and the relationship
between the families in the Bonnetiaceae, Theaceae and Guttiferae, and G. van Vliet the
identification and systematic and phylogenetic relationships within the Rhizophoraceae. R.
den Outer and W. van Veenendaal investigate the effects of the rainforest and savanna
environment (in West Africa) on within-species wood anatomical characters and try to
distinguish them from the effects of phylogenetic drift. A short paper contributed by D. F.
Cutler deals with a little-known field, variability of root wood structure and its comparison
with trunk wood.
_, There have been many publications in recent years which deal with the distribution of
siliceous inclusions and their value in taxonomy. In the present volume Ben ter Welle gives
a comprehensive survey of the occurrence of silica in neotropical woods and concludes that
it has considerable diagnostic value, especially at the species level. Probably the most
interesting and significant contribution in this book is that of P. Baas, on xylem vessel
variation in relation to environment and evolution. He gives a careful and well documented
cane of Carlquist’s theory of adaptive diversification, for which he finds rather limited
evidence.
Book Reviews 315
The book includes some interesting chapters on more general topics, such as aspects of
branch abscission (J. Koek-Noorman and B. ter Welle), a somewhat neglected subject, and
the examination of a theoretical model for liquid flow through tracheids and pits (A. J.
Bolton). The cambial production of wood in relation to apical meristem activity is explored
by M. P. Denne, and the origins of short term tracheid variation by E. D. Ford and A. W.
Robards. Only three papers are of essentially technological interest, two dealing with
structural failure and one with plastic impregnation, in relation to wood microstructure.
This book is exceptionally well produced. There is an attractive layout and good
reproduction of the line drawings and photomicrographs. A particularly valuable feature is
the adequate lists of references, to literature much of which may not be familiar to every
botanist.
A. R. A. NOEL
Preliminary Announcement
THIRTEENTH INTERNATIONAL BOTANICAL CONGRESS
Sydney, Australia. 21—-28th August, 1981
The Programme will consist of 12 sections—molecular, metabolic, cellular and structural,
developmental, environmental, community, genetic, systematic and evolutionary, fungal,
aquatic, historical, and applied botany. There will be plenary sessions, symposia, and
sessions for submitted contributions (papers and posters). Chairman of the Programme
Committee: Dr L. T. Evans.
Field Trips will include visits to arid and semi-arid regions, eucalypt forest, rain forest,
heath, coastal vegetation (e.g. Great Barrier Reef, mangroves) etc., and specialist trips.
Chairman of the Field Trips Committee: Prof. L. D. Pryor.
First Circular, containing details, will be mailed in August, 1979. Send your name and full
address, preferably on a postcard, to ensure your inclusion on the mailing list.
Enquiries should be sent to the Executive Secretary, Dr W. J. Cram.
Congress address—13th I.B.C., University of Sydney, N.S.W. 2006, Australia.
Sponsored by the Australian Academy of Science
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JIS. Afr. Bot. 44 (4): 317-328 (1978)
THE VASCULAR ANATOMY OF THE SEEDLING OF ENCEPHALARTOS
EUGENE-MARAISIT VERDOORN
P. J. ROBBERTSE, S. VAN DER WESTHUIZEN AND G. K. THERON
(Department of Botany, University of Pretoria)
ABSTRACT
The path of the vascular strands in the seedling of Encephalartos eugene-maraisii, from
the primary root to the cotyledons and first vegetative leaves is discussed. Each of the two
cotyledons is supplied with three leaf traces while the first vegetative leaf is supplied with
four vascular bundles orginating from a girdle in the vascular plate. The primary xylem in
the leaf base is mesarch, becoming endarch towards the first pinna.
UITTREKSEL
DIE VAATANATOMIE VAN ENCEPHALARTOS EUGENE-MARAISII VERDOORN
SAAILINGE
Die ligging van die vaatbundels in die Encephalartos eugene-maraisii saailinge word
vanaf die primére wortelpunt tot by die saadlobbe en eerste vegetatiewe blare bespreek.
Elk van die twee saadlobbe is voorsien van drie blaarvaatstringe terwyl die eerste vegeta-
tiewe blare van vier vaatstringe wat uit ’n gordel in die vaatplaat ontstaan. Die primére
xileem in die blaarbasis is mesarg en word endarg in die rigting van die eerste pinna.
INTRODUCTION
The Cycadales is a very interesting plant group and it is not surprising that by
the end of the 19th century a considerable amount of literature had been built up
on a variety of morphological aspects of this taxon. Very little, however, has been
done on the South African Encephalartos species and therefore a morphological
study of E. eugene-maraisii was started by van der Westhuizen (1976). During
this study it became clear that there was still doubt as to the origin of the
centrifugal xylem in the petiole of the Cycads in spite of Le Goc’s ( 1914) paper
which dealt with this problem. There was also doubt about the number of vascular
strands entering the cotyledons of seedlings of Encephalartos species since Matte,
according to Dorety (1908), described the anatomy of E. barteri with three
cotyledons (probably an abnormal seedling), each with only one trace.
This study was undertaken to contribute to the knowledge of the anatomy of
the Cycadales and to try to supply answers to the above-mentioned problems.
MATERIAL AND METHODS
Two-month-old seedlings of Encephalartos eugene-maraisil and of E. trans-
venosus were obtained from the Division of Nature Conservation, Transvaal
Accepted for publication 25th November, 1977.
SY)
318 Journal of South African Botany
Provincial Administration. One seedling was transversely sectioned at the posi-
tions indicated in Fig. 1. Diagrams of the sections are shown in Fig. 2A-S. Serial
microtome sections, 8—10 jm thick, were made of one other seedling and hand
sections were made of two other seedlings. Standard procedures for embedding,
sectioning and staining were followed to obtain serial sections (Johansen, 1940).
RESULTS AND DISCUSSION
At the seedling stage depicted in Fig. 1, one vegetative leaf has developed
while the greater part of the two cotyledons is still embedded in the female
gametophyte tissue which is enveloped by the seed coat. The proximal end of the
primary root is tuberous, and a few young apogeotropic roots have developed (not
shown in figure).
At levels A and B (Figs | and 2) the roots of some seedlings are octarch while
others are tri- or tetrarch. In the root from which the sections A—S were prepared
no endodermis was distinguishable although it was well-developed below this
point in all the roots examined. The pith is well-developed and its diameter
increases in the tuberous part of the root, due to irregular divisions of cells of the
ground tissue, resulting in the displacement of the primary xylem and phloem
groups (Fig. 2B—D). This primary thickening growth also takes place in the cortex.
The primary xylem groups consist of relatively small protoxylem elements and
large metaxylem elements which give the impression of transfusion tissue. These
metaxylem elements do not only differentiate in a centripetal direction, but also in
a lateral and centrifugal direction to form arcs radiating from the protoxylem.
Worsdell (1897) also commented on the extension of the centripetal xylem
towards the sides of the bundle in the cotyledonary bundles of Cycas and Ginkgo,
in his attempts to determine the origin of the transfusion tissue in Gymnosperm
leaves.
The later-formed metaxylem elements at first lie in direct contact with the
other metaxylem elements but can at a later stage become separated from them by
the intrusion and divisions of the pith parenchyma cells (Fig. 5A).
In the seedling with the octarch root (Fig. 2A) xylem strands 1 and 7
dichotomise at levels C and D respectively, but branch la disappears again at level
D and branch 7a disappears at level E. At level F, branches 1b and 7b fuse while 3
and 6 disappear so that at the lower end of the hypocotyl at level G only branches
2, 4, 5 and 7b remain.
In tetrarch roots, the primary xylem strands remain unchanged up to level G,
while in the case of a triarch root, one of the xylem strands dichotomises to form a
tetrarch stele in the hypocotyl. Pearson (1898) describes a reduction of the
pentarch structure to a triarch in Bowenia while Dorety (1919), in her description
of the seedling of Dioon spinolosum, states that ‘the vascular cylinder of the
hypocotyl is a protostele; it has four easily recognized protoxylem groups, in no
way differing from the hypocotyl cylinders of Ceratozamia and Microcycas’’.
Vascular anatomy of seedling of Encephalartos eugene-maraisii Verd. 319
BGI
Young seedling of Encephalartos eugene-maraisii showing levels A-B where sections were
made. cot—cotyledon; sc—seed coat; vi—vegetative leaf.
About the seedling of Dioon edule, Thiessen (1908) states that at each corner of
the squarish vascular plate there is a group of protoxylem elements. In E.
transvenosus the vascular cylinder is pseudotetrarch. It seems, therefore, that
regardless of the number of protoxylem strands in the roots of most of the Cycad
320 Journal of South African Botany
seedlings investigated, the hypocotyl is tetrarch or triarch. The triarch condition in
Bowenia will have to be re-investigated as the condition in E. eugene-maraisii at
level H (Fig. 2), which resembles that in Bowenia to a certain extent, could be
misinterpreted as triarch.
The hypocotyl is very short and stretches only from levels G to about K, a
length of approximately one millimeter. This is in agreement with what has been
found in other Cycad seedlings.
Each of the two cotyledons is supplied with three leaf traces. Traces a, b and c
(Fig. 2L) enter the one cotyledon while traces d, e and f (Fig. 2M) enter the
second (see also Fig. 3). The central traces b and e remain unbranched while the
lateral traces c, d and f dichotomise and a splits up into three branches (Fig.
2L—O). In the other seedlings of E. eugene-maraisii investigated, trace a splits up
into two branches.
Three leaf traces (strands) were also found in Cycas revoluta (Worsdell, 1898),
Cycas siamensis (Matte, 1904), Ceratozamia (Dorety, 1908), Macrozamia spiralis
(Worsdell, 1898) and Stangeria paradoxa (Worsdell, 1898) while in Dioon edule
(Thiessen, 1908) and Dioon spinolosum (Dorety, 1919) two leaf traces are given
off to each cotyledon but, before entering the cotyledons, they branch so that four
vascular strands enter each cotyledon. In Zamia muricata (Karsten, 1856) and
Bowenia spectabilis (Pearson, 1898) only one leaf trace per cotyledon is given off
but it also branches so that each cotyledon is supplied by four vascular strands.
According to Coulter & Chamberlain (1910) the condition in Dioon is characteris-
tic of the Cycads but from the cited examples it is clear that the three leaf traces
per cotyledon as described in this paper for Encephalartos is more common
among the Cycads. It would seem, therefore, that the seedling of E. barteri with
three cotyledons which was described by Matte (1904) must have been abnormal.
Three leaf traces were also found in a seedling of E. transvenosus which was
studied as a comparison with seedlings of E. eugene-maraisii (Fig. 4).
In his description of the seedlings of Cycas revoluta, Macrozamia spiralis and
Stangeria paradoxa, Worsdell (1898) mentioned three peculiar strands leaving the
stele in the hypocotyl and extending ‘‘perpendicularly’’ in the cortical tissue. He
could not decide whether they were strands of roots or not. In this study similar
Strands were observed but they were definitely the strands of the so-called
apogeotropical roots which later dichotomised to form coralloid roots (Fig. 4, see
also Fig. 5D for gaps where these strands leave the central cylinder).
Four vascular bundles enter the first vegetative leaf at two different levels.
Bundles h and i which remain unbranched, have their origin at level O (Fig. 2).
Both are compound bundles receiving traces from a common vascular arc between
two of the original vascular strands of the hypocotyl. Bundles g and j originate at
level M on the side of the leaf base opposite to the root poles 2 and 4 (Fig. 3)
forming a girdle around the vascular plate. The bundles g and j each divide into
four bundles which enter the leaf base at levels P and Q (Fig. 2C). The first
Vascular anatomy of seedling of Encephalartos eugene-maraisii Verd. 321
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Serial transverse sections through a seedling of E. eugene-maraisti at le :
Fig. 1. 1-8: primary xylem Erope in root and hypocotyl. a-f: vascular strands Supp ne
the cotyledons. g—j: vascular strands supplying the first vegetative ee aa ia
meristem; cot 1 and cot 2—basis of cee Beg boa ne pa a ee
second hloem; sp—secondary parenchyma, sx—secon 5 it
pelle oP rasculas plate: fenreet vegetative leaf; 2ndl—second vegetative leaf.
B22 Journal of South African Botany
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Serial transverse sections through a seedling of E. eugene-maraisii at levels indicated in
Fig. 1. 1-8: primary xylem groups in root and hypocotyl. a-f: vascular strands supplying
the cotyledons. g—j: vascular strands supplying the first. vegetative leaf. am—apical
meristem; cot 1 and cot 2—basis of cotyledons; per—periderm; sd—slime ducts; sf—
secondary phloem; sp—secondary parenchyma; sx—secondary xylem; t—tanniniferous
cells; vpl—vascular plate; 1stl—first vegetative leaf; 2ndl—second vegetative leaf.
Vascular anatomy of seedling of Encephalartos eugene-maraisii Verd. 323
Fic. 2
Serial transverse sections through a seedling of E. eugene-maraisii at levels indicated in
Fig. 1. 1-8: primary xylem groups in root and hypocotyl. a—f: vascular strands supplying
the cotyledons. g—j: vascular strands supplying the first vegetative leaf. am—apical
meristem; cot 1 and cot 2—basis of cotyledons; per—periderm; sd—slime ducts; sf—
secondary phloem; sp—secondary parenchyma; sx—secondary xylem; t—tanniniferous
cells; vpl—vascular plate; 1stl—first vegetative leaf; 2ndl—second vegetative leaf.
324 Journal of South African Botany
Fic. 3.
Reconstruction of sections A—S in Fig. 2 to illustrate the vasculature in the seedling of E.
eugene-maraisii. 2, 4, 5 & 7: vascular groups of tetrarch stele. a—f: vascular strands
supplying the cotyledons. g—j: vascular strands supplying the first vegetative leaf. x: girdle
of vascular tissue supplying the 2nd vegetative leaf.
vegetative leaf thus has a total of ten vascular strands in its base (Fig. 2CR). In E.
transvenosus (Fig. 4) the same basic pattern is followed.
In his description of the vascular anatomy of the seedling of Dioon edule,
Thiessen (1908) found four vascular bundles entering the first vegetative leaf. A
similar condition occurs in E. eugene-maraisii with the exception that in E.
eugene-maraisii the first two bundles are compound, whereas in Dioon they are
not. Dorety (1908) also found a similar four-bundle pattern in Ceratozamia.
Secondary growth in the root commences when cambium segments are
Vascular anatomy of seedling of Encephalartos eugene-maraisii Verd. 325
Fic. 4.
Reconstruction of the vasculature in the seedling of E. transvenosus. Figures and numbers
as for Fig. 3. ag—apogeotropic roots.
differentiated on either side of each radiating primary xylem strand, thus forming
secondary tracheids and sieve cells in a more or less tangential direction (Figs 2F
and 5C). Because secondary tracheids are about the same size and have the same
Staining qualities as the metaxylem elements, it is very difficult to differentiate
between them. Opposite the protoxylem, a few secondary tracheids may be formed
although most of the secondary tissue at this point consists of secondary paren-
chyma, in broad medullary rays (Fig. 5B).
Le Goc (1914) pointed out that the parenchymatous band between the protoxy-
lem and centrifugal xylem in the petiole of Cycadales is an indication that the
centrifugal xylem is of secondary origin. In this study, however, it has been found
that even in the root, the metaxylem elements are often separated from the
protoxylem by the intrusion of parenchyma cells as a result of primary thickening
growth of the tuberous roots (Fig. 5A). The presence of the parenchymatous band,
therefore does not provide sufficient evidence for making deductions as to the
origin of the centrifugal xylem.
326 Journal of South African Botany
Transverse sections at different levels of the seedling of E. eugene-maraisii. A at level B in
Fig. 2; B at level F in Fig. 2; C at level D in Fig. 2; D from another seedling at level H.
mc—meristematic cortical cell; mr—medullary ray parenchyma; mx—metaxylem tracheids,
px—protoxylem; sx and xs—secondary xylem tracheids; to—trace gaps for apogeotropical
roots; vc—vascular cambium.
When the differentiation of xylem elements from the procambium strands in
the first vegetative leaf is followed acropetally from the leaf base into the petiole,
it is clear that the position of the protoxylem in the procambium strand in these
two regions differs. In the leaf base the first protoxylem elements differentiate near
the centre of the procambium strand (Fig. 6A), while the metaxylem elements
differentiate centrifugally as well as centripetally, thus giving rise to mesarch
xylem (Fig. 6C). Acropetally the first protoxylem elements are gradually displaced
centrifugally so that less centrifugal and more centripetal xylem is formed until,
near the first pinna, no centrifugal xylem differentiates at all, thus giving rise to
Vascular anatomy of seedling of Encephalartos eugene-maraisii Verd. 327
Fic. 6
Segments of transverse sections of the young petiole of E. eugene-maraisii to show
A—procambium strand with first protoxylem elements in the leaf base; B—exarch xylem
near first pinna; C—mesarch xylem in middle of petiole. px—protoxylem elements;
pp—primary phloem; ex—exarch xylem group; me—mesarch xylem group.
exarch xylem (Fig. 6B). After secondary thickening growth has commenced,
therefore, the centrifugal xylem in the leaf base would consist of metaxylem and
secondary xylem whereas in the distal end of the petiole it would consist purely of
secondary xylem.
Le Goc (1914) stated that ‘‘. . . the centripetal xylem is an independent tissue,
probably the remnant of an ancient and more developed structure’. He also stated
that the centripetal and centrifugal xylem ‘‘. . . are continuous physiologically but
not morphologically’’, probably having the parenchyma cells between the protoxy-
328 Journal of South African Botany
lem and the centrifugal xylem in mind. If this was true it would mean the
centrifugal xylem would consist only of metaxylem and it would be hard to
believe that it could be an ‘‘independent tissue’.
It is therefore concluded that:
1. The primary centripetal and centrifugal xylem has a common group of
protoxylem elements and therefore cannot be regarded as independent tissues.
Instead it is found in vascular bundles with mesarch primary xylem.
2. Before secondary growth has commenced, the centrifugal xylem consists only
of metaxylem elements but it can be supplemented by secondary xylem once
secondary thickening growth has taken place.
ACKNOWLEDGEMENTS
This research was sponsored by the Division of Nature Conservation of the
Transvaal Provincial Administration and the University of Pretoria. Thanks are
due to Mrs J. E. Lombard for reading the manuscript.
REFERENCES
COULTER, J. M. and CHAMBERLAIN, C. J., 1910. Morphology of the Gymnosperms.
Chicago: University of Chicago Press.
Dorety, H. A., 1908. The seedling of Ceratozamia. Bot. Gaz. 46: 203-220.
Dorety, H. A., 1919. Embryo and seedling of Dioon spinulosum. Bot. Gaz. 67:
25-257.
JOHANSEN, D. A., 1940. Plant microtechnique. New York: McGraw-Hill.
KaRSTEN, G., 1856. Organographische Betrachtung der Zamia muricata Willd.
Abh. Kgl. Akad. Wiss. Berlin. 193-219.
Le Goc, M. J., 1914. Observations on the centripetal and centrifugal xylems in
petioles of cycads. Ann. Bot. 28: 184-193.
Matte, H., 1904. Recherches sur l'appareil libéro-ligneux des Cycadacées.
Caen.
PEARSON, H. H. W., 1898. Anatomy of the seedling of Bowenia spectabilis Hook. f.
Ann. Bot. 12: 475-490.
THIESSEN, R., 1908. The vascular anatomy of the seedling of Dioon edule. Bot.
Gaz. 46: 357-380.
VAN DER WESTHUIZEN, S., 1976. ’n Morfologiese studie van Encephalartos
eugene-maraisti Verdoorn. M.Sc. thesis. Pretoria: University of Pretoria.
WorsbeELL, W. C., 1897. On the origin of transfusion tissue in leaves of
Gymnospermous plants. J. Linn. Soc. 33: 118-122.
WorspELL, W. C., 1898. Comparative anatomy of the Cycadaceae. J. Linn. Soc.
33: 437-457.
|
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So a een oa
JIS. Afr. Bot. 44 (4): 329-362 (1978)
STUDIES IN THE GENERA OF THE DIOSMEAE (RUTACEAE): 7.
ION WILLIAMS
(Honorary Reader, Bolus Herbarium, University of Cape Town
Honorary Research Worker, Compton Herbarium, Kirstenbosch)
ABSTRACT
Acmadenia laevigata Bartl. & Wendl. is described afresh as Euchaetis meridionalis.
Euchaetis laevigata Turcz. (= Acmadenia assimilis Sond.) is redescribed in order to clarify
its difference from Euchaetis meridionalis. Two new species of Euchaetis, four new
species of Acmadenia and one new species of Macrostylis are described. Diosma wit-
tebergensis is transferred to the genus Acmadenia.
UITTREKSEL
STUDIES IN THE GENERA VAN DIE DIOSMEAE (RUTACEAE): 7
Acmadenia laevigata Bartl. & Wendl. word as Euchaetis meridionalis opnuut beskryf.
Euchaetis laevigata Turcz. (= Acmadenia assimilis Sond.) word weer beskryf om sy
verskil van Euchatis meridionalis te beklemtoon. Twee nuwe Euchaetis soorte, vier nuwe
Acmadenia soorte en een nuwe Macrostylis soort word beskryf. Diosma wittebergensis
word by Acmadenia ingedeel.
Euchaetis meridionalis Williams, sp.nov. affinis E. scabricosta Williams sed
costis paucioribus glanduloso-punctatis, foliis orbicularibus non lanceolatis, et
capitulis ad basim in axilibus resiniferis differt.
Frutex ad 1,5 m, ad basim monocaulis. Rami numerosi, erecti, glabri, brevi.
Ramuli numerosi, erecti, brevi, sparsim puberuli, dense foliosi. Folia ad 6 mm
longa, 3,5 mm lata, elliptica vel orbiculares, obtusa, 4-5 faria, sub-complicata, breve
petiolata, sub-imbricata, erecta, glabra, sparsim ciliolata, costis marginibusque
pauci-glanduloso-punctatis. Inflorescentia terminilia, 5—6 nata. Bractea 4 mm
longa, 2—3 mm lata, elliptia vel orbiculares, obtusa, glabra, sessilia, crassiuscula,
intus ad basim resiniferis, marginibus anguste cartilagineis sparsim spinulosis.
Bracteolae duae, 2,5 mm longae, 1,2 mm latae, sub-ellipticae, glabrae, ciliolatae,
ad basim resiniferis. Sepala quinque, 2,5 mm longa, 2—2,3 mm lata, sub-elliptica,
glabra, sparsim ciliolata. Petala quinque, 5,2 mm longa, 3,1 mm lata, sub-rosea
vel alba, orbiculata, unguibus transverse barbatis, ciliatis. Staminodia quinque,
vestigiala. Fila quinque, glabra, post anthesin 1,5 mm longa. Antherae quinque,
1,2 mm longae, 0,9 mm latae, roseae, apicibus minute glandulosis. Pollen 45-50
pu longum, 30-33 p latum, oblongum. Discus ovarium excedens, viridus, nec-
tarifer. Stigma 0,5 mm diam., capitatum. Stylus 1,4 mm longus, glaber, erectus.
Ovarium 5-carpellatum, 1 mm diam., 0,9 mm longum, apicibus obtusis sparsim
Accepted for publication 20th January, 1978.
329
330 Journal of South African Botany
setulosis. Fructus 5-carpellatus, 7 mm longus, 4,5 mm diam., glaber, multi-
glandulosis, cornibus 2 mm longis, erectis. Semen 3,8—4,2 mm longum, 1,5 mm
latum, piceum, nitens.
Type: CAPE—3420 (Bredasdorp): De Hoop behind sand dunes on track to Koppie
Alleen, Bredasdorp Division (-AD), 14/4/1972, Williams 1644 (NBG, holotype).
Acmadenia laevigata Bartl. & Wendl. (1824) is the earliest name that was
applied to this species but the epithet “‘laevigata’’ cannot be transferred to
Euchaetis due to the prior existence of the name Euchaetis laevigata Turcz.
(1858). The name Euchaetis meridionalis is therefore proposed to embrace this
taxon which is here described de novo.
Shrubs up to 1,5 m, usually much less, forming a dense rounded bush, single
stemmed at base, sometimes in close mats. Branches numerous, erect, short,
glabrous, di-trichotomous, somewhat rough with leaf scars. Branchlets numerous,
erect, short, sparsely puberulous, densely foliate. Leaves up to 6 mm long, 3,5
mm. broad, elliptic or orbicular, obtuse, sub-imbricate, erect, glabrous, 4—5
ranked, somewhat complicate; midrib prominent with 2 or 3 gland dots; margins
narrowly cartilaginous, sparsely ciliolate with a few gland dots; petiole short.
Inflorescence terminal, 4—6 nate, flowers pale pink to white. Bract leaf-like, 4 mm
long, 2-3 mm broad, elliptic or orbicular, obtuse, glabrous, sessile, somewhat
thick, exuding resin from numerous glands situated towards the axil; margins
narrowly cartilaginous, sparsely ciliolate. Bracteoles two, 2,5 mm long, 1,2 mm
broad, asymmetrically elliptic, glabrous, ciliolate, apex thickened and green,
resiniferous below. Calyx lobes five, 2,5 mm long, 2—2,3 mm broad, sub-elliptic,
glabrous, pinkish; apex thickened; margins broadly translucent below, sparsely
ciliolate. Petals five, 5,2 mm long; blade 2,2 mm long, 3,1 mm broad, glabrous,
pale pink to white; claw 3 mm long, 1,8 mm broad, transversely bearded and
sparsely ciliate above. Staminodes five, vestigial or a stalked gland 0,2 mm long.
Filaments five, becoming 1,5 mm long, glabrous, pale pink, acicular. Anthers five,
1,2 mm long, 0,9 mm broad, pink, apex with a minute gland. Pollen 45-50 wu
long, 30-33 w broad, oblong. Disc dark green, exudes nectar, curling inwards
over the ovary. Stigma 0,5 mm diam., globose, dark green, capitate. Style
becoming 1,4 mm long, glabrous, erect. Ovary 5-carpellate, 1 mm diam., 0,9 mm
long; /obes obtuse with a few hairs towards the sides above. Fruit 5-carpellate, 7
mm long, 4,5 mm diam., surrounded for the most part by the persistent calyx and
petals; carpel glabrous with many gland dots; horn 2 mm long, erect with a few
hairs above, apex retuse with a gland. Seed 3,8—4,2 mm long including the white
aril, 1,5 mm broad, black, shining.
SPECIMENS EXAMINED
CAPE—3420 (Bredasdorp): De Hoop, behind sand dunes on track to Koppie
Alleen (-AD), 200 ft., 14/4/1972, Williams 1644 (NBG); De Hoop, flats, lime-
Studies in genera of Diosmeae (Rutaceae): 7 331
stone hills and along pass to Wydgelegen, 8/4/1957, Lewis 5130 (BOL, NBG),
9/6/1970, Esterhuysen 32461 (BOL), 10/3/1957, Barker 8735 (NBG), —/7/1962,
Macpherson s.n. (NBG), 5/2/1963, Jordaan M227 (STE), c 100 ft., 28/7/1970,
v.d. Merwe 137 (STE); De Hoop, N. side of pass, 415 ft., 9/4/1974, Williams
1888 (NBG); De Hoop, S. side of pass, 200 ft., 9/4/1974, Williams 1890 (NBG),
175 ft., 27/11/1974, Williams 1940 (NBG): Potberg, Elands Pad, near the coast
(-BC), 100 ft., 12/4/1972, Williams 1638 (NBG); Cape Infanta, Sebastian Bay
(-BD), 28/9/1959, Esterhuysen 28314 (BOL), 23/9/1947, Walgate 874 (NBG),
Kunth s.n. (MEL 52824); Arniston, Wagenhuiskraal, Kliprug (-CA), 15/9/1944,
Henrici 3716 (BOL, NBG), 5/6/1900, Fry s.n. (GRA, PRE), 0-150 ft.
24/8/1962, Taylor 3807 (PRE, STE), 90 ft., 20/9/1968, Marsh 919 (PRE, STE),
—/7/1933, Jordaan s.n. (STE 18672); Dronkvlei, 100 ft., 26/8/1963 van Breda
1636 (K, PRE); 3,4 miles N.W. of Skipskop (-CB), 100 ft., 22/6/1962, Acocks
22268 (K, PRE); Skipskop behind the dunes, 50 ft., 9/4/1974, Williams 1891
(NBG); Struis Bay near sea (-CC), 1/12/1933, Salter 4104 (BM, BOL, K), 50 ft.,
21/6/1972, Williams 1661 (NBG); Cape Agulhas 2 miles east and rocks along
coast, 16/9/1934, Salter 4836 (BM, BOL, K), 5/8/1940, Esterhuysen 2975
(BOL, GRA, K, NBG, SAM), 4/8/1940, Compton 9081 (NBG); prope Zoutendals
valley, —/10/—, Ecklon & Zeyher 824(S).
Without locality: Scholl s.n. (B-W 4784 as to r.-hand specimen), Mundt s.n.
(SAM 30579 as to r.-hand specimen), Guenzius s.n. (W), Sieber s.n. (W).
(Ecklon & Zeyher 826 (S, SAM) from between Pampoenkraal and Paarl is an
extremely doubtful locality).
DISTRIBUTION AND BIOLOGY
Euchaetis meridionalis occurs in a fairly narrow coastal area lying between
Cape Agulhas and Cape Infanta usually close to the coast but, at only one point
near Wydgelegen, reaching as far as thirteen kilometres from the sea. It is by no
means rare and can form dense mats in exposed areas near to the sea (Esterhuysen
2975). It is usually found at altitudes of from 15 to 60 m (50-200 ft.) above sea
level, growing on limestone associated with the Bredasdorp geological series. It
has been observed in bloom from April to December and ripe fruits have been
collected from August to November. The small but conspicuous flowers, the sticky
pollen and the presence of nectar indicate that this plant is most probably
pollinated by insects. An assumption confirmed by the observation of honey bees
visiting the flowers (Williams 1 638). Regeneration takes place, probably only after
fires, from seed which has been ejected when ripe from the capsule by the usual
catapult mechanism. The leaves, when crushed, have an odour reminiscent of
cedar and may exhibit a whitish waxy exudate originating as a sticky resin near
the axils of the young leaves. On farm lands the plants have been observed to have
been grazed by stock (Williams 1644, Henrici 3716).
332 Journal of South African Botany
VARIATION
It is quite noticeable that the leaves of plants from near the sea at Struisbaai
are shorter and broader than those of the plants from further inland and that those
from the farthest inland, near to Wydgelegen, are beginning to look very much
like those of Euchaetis scabricosta indicating a very close relationship between
these two species.
DISCUSSION
Because of the transversely bearded petals this plant may be regarded as a
typical Euchaetis. It may be distinguished from Euchaetis scabricosta, as previ-
ously mentioned (Williams, 1974), having a midrib with fewer gland dots, leaves
that are more obtuse, more crowded, shorter and broader, with a resinous exudate
towards the leaf axils. Furthermore a recent gathering of Euchaetis scabricosta
(Williams 2181) from near Bredasdorp has shown that, whereas E. scabricosta
may regenerate after fires by sprouting from the rootstock just below ground level,
E. meridionalis, being single stemmed at base, has never been observed to do this.
A most notable difference between these two species is that E. meridionalis is
only found on limestone whereas E. scabricosta is found in sandy soil.
REFERENCE
WILLIAMS, Ion, 1974. JIS. Afr. Bot. 40 (4): 286-289.
; i Fic. 1.
Euchaetis meridionalis: 1, leaf. 2, bracteole. 3, bract. 4, calyx lobe. 5, petal. 6, anther. 7,
gynoecium. 8, pollen. 9, distribution.
Studies in genera of Diosmeae (Rutaceae): 7 333
Euchaetis laevigata Turcz. in Bull. Soc. Imp. Nat. Mosc. 31. 1-438 (1858).
Type: between Cape Agulhas and Potberg, on limestone hills, under 500 tite
3/8/1831, Drege IV, C, a, 2 (BM, E, K, MEL, P, S, isotypes).
Acmadenia laevigata E. Mey in Drege Zwei pflanzen. Doc.: 122, 161 (1844).
nom. nud.
Acmadenia assimilis Sonder in Flor. Cap. 1: 383 (1860). Type: as above.
It is most extraordinary to think that this plant collected by Drége in 1831 on
limestone hills between Cape Agulhas and Potberg should have passed unnoticed
until comparatively recently when Acocks gathered specimens on the north side of
the limestone hills between Bredasdorp and Wydgelegen. Following Acocks’
lead, further material was collected from quite near Bredasdorp, just beyond the
lime quarry on the Swellendam road. It was found to be a perfect match with the
specimens collected by Drege and named Acmadenia laevigata by E. Meyer.
Sonder, unaware of Turcszaninov’s paper, named Drege’s plant Acmadenia
assimilis, the name Acmadenia laevigata having been previously used by Bartling
and Wendland for another taxon in this genus. Although Sonder’s action was
correct, local botanists were confused and for many years assigned plants found
between Cape Agulhas and Potberg to Acmadenia assimilis in error. Because of
this misidentification it has been deemed advisable to redescribe Euchaetis
laevigata at the same time as publishing the description of Euchaetis meridionalis
which now replaces Acmadenia laevigata B. & W. At the same time attention is
drawn to the fact that the three species mentioned by Sonder: (11) A. laevigata B.
& W., (12) A. pungens B. & W. and (13) A. assimilis Sond., considered by
Sonder to belong in Acmadenia, all possess transversely bearded petals and
rudimentary staminodes and have therefore been placed in Euchaetis under the
names E. meridionalis Williams, E. pungens (B. & W.) Williams and E.
laevigata Turcz. in that order.
DESCRIPTION
Shrubs up to 0,8 m tall, erect, forming a dense bush single stemmed at base.
Branches erect, often dichotomous, slender, with a greyish bark becoming fairly
smooth and leafless after about three seasons. Branchlets numerous, erect, slender,
somewhat reddened, glabrous, except for some minute puberulence, often in
opposite pairs. Leaves 2,8-3,6 mm diam., orbicular, sessile, 4-ranked, opposite,
recurved-erect, glabrous, very sparsely and minutely ciliolate; apex with a blunt
point; margins narrowly translucent and somewhat denticulate, subcomplicate with
a distinct midrib and scattered oil glands; adaxial surface somewhat concave.
Inflorescence terminal, with the white flowers in opposite pairs or fours below
which the next years shoots emerge. Bract one to each flower, 2,5 mm long
1,6-2,4 mm broad, ciliolate, leaf-like. Bracteoles two 2,5 mm long, 1,3-1,6 mm
broad, elliptic, obtuse, glabrous, keeled, apex thickened; margins minutely cilio-
late and translucent below. Calyx lobes five, about 2,5 mm diam., asymmetrically
334 Journal of South African Botany
orbicular, glabrous, minutely ciliolate, thickened and spreading at the apex; the
margins broadly translucent below. Petals five, 4,5 mm long; limb 1,6-1,8 mm
long, 2,2—2,5 mm broad, sub-orbicular, glabrous, pinkish-white; claw 2,5 mm
long, 1,5 mm broad narrowing to 0,8 mm below, more or less translucent,
transversely bearded above, ciliate, a few hairs down the prominent midrib.
Staminodes five, vestigial, a minute gland about 0,1 mm long at most, situated low
down on the outside of the disc. Filaments five, at first 0,5 mm long becoming 1,8
mm long after anthesis and placing the discharged anther outside the perimeter of
the claws. Anthers five, 1 mm long before anthesis, reddish-orange, with a minute
apical gland. Pollen 50-55 pw long, 28-30 mw broad, oblong. Disc minutely
punctate, dark green, closes somewhat over the ovary, exudes nectar. Stigma 0,6
mm diam., globose, capitate. Style becoming 1,2 mm long, glabrous, erect,
persisting. Ovary 5-carpellate, each carpel with a gland clasped at the apex and
minute spiky lumps along adjacent edges. Fruit 5-carpellate, 5,5 mm long, about
3,8 mm diam., glabrous, smooth; carpels with about 6 gland dots along the
margins and a few very short hairs above; horns 1 mm long or less, erect, clasping
a small gland at the apex, equalling the persistent stigma. Seed (from Williams
2025) 2,7—3 mm long with the white aril produced a further 0,6 mm, 1,5 mm
broad, black, shining.
SPECIMENS EXAMINED
CAPE—3420 (Bredasdorp): van der Stelskraal (-AC), 400 ft., 1/8/1968,
Acocks 24042 (PRE), 300 ft., 2/7/1972, Williams 1662 (NBG); Soutpansvlakte, 8
miles E. of Bredasdorp, 340 ft., 4/6/1974, Williams 1904 (NBG); 3 km N.E. of
Bredasdorp on road to Swellendam (-CA), 300 ft., 21/11/1973, Williams 1874
(NBG), 3/8/1975, Williams 2025 (NBG); between Cape Agulhas and Potberg, on
limestone hills, under 500 ft., 3/8/1831, Drége IV, C, a, 2 (BM, E, K, MEL, P, S,
isotypes).
DISTRIBUTION AND BIOLOGY
Euchaetis laevigata occurs quite frequently over a rather small area about 23
km long on the N side of the limestone ridge running NE from Bredasdorp at an
altitude of 90-120 m (300-400 ft.) above sea level. It was found in full bloom in
the middle of October but at other times only a few plants with flowers were seen
and only one plant with fruits was found in August. It is possible that the
extremely dry winters experienced at that time may have been the reason. The
plant is single stemmed at base and apparently will not regenerate after fires from
the stump so that one presumes that it can only regenerate from seed.
VARIATION
No variation or hybridisation had been observed but the plants at van der
Stelskraal were the tallest and most healthy.
Studies in genera of Diosmeae (Rutaceae): 7 33/5
DISCUSSION
The transversely bearded petals and the vestigial staminodes indicate that this
plant is a typical Euchaetis.
It differs from both Euchaetis scabricosta and Euchaetis meridionalis in
having leaves that are always Opposite with the midrib recurved, somewhat
smaller, orbicular in outline with gland dots scattered and not just on midrib and
margins. The shoots are dry without any trace of resin or wax.
50u
Fic. 2.
Euchaetis laevigata: 1, leaf. 2, bract. 3, bracteole. 4, calyx lobe. 5, petal. 6, anther. 7,
staminode. 8, pollen. 9, distribution.
Euchaetis intonsa Williams, sp. nov., propria propter folia alterna elliptica
sparsim pubescentia ciliata, bracteae bracteolae et sepala pubescentes, petala parva
costis intus glabris.
Frutex 400 mm, ad basim monocaulis. Rami numerosi, erecti, glabrescenti,
cinerei. Ramuli numerosi, erecti, puberuli, graciles, ex parte rubelli, foliosi,
irregulariter alterni. Folia 4 mm longa, 2,2—2,4 mm lata, elliptica, sub-acuta,
sub-erecta, pleurumque alterna; costis prominentibus, glanduloso-punctatis, spar-
sim pubescentibus, ad apex callosis; marginibus sparsim ciliatis, minute
glanduloso-punctatis. /nflorescentia terminalis, sessilis, 3—4 nata. Bractea una,
2,7 mm longa, 1,1—1,7 mm lata, oblanceolata, obtusa, ciliata, sparsim pubescens.
Bracteolae duae, 2,5 mm longae, 1 mm latae, oblongae, obtusae, ciliatae,
pubescentes. Sepala quinque, 2,6 mm longa, 2 mm lata, asymmetrice oblonga,
conspicua, purpurata, ciliata, sparsim pubescentia. Petala quinque, 3,8—4,2 mm
longa; limbus 1,3—1,8 mm latus, sub-orbicularis, subroseus; unguis 0 2-1 mm
336 Journal of South African Botany
latus, forte transverse barbata, ciliata. Staminodia quinque, 0,1 mm longa, ves-
tigiala. Filamenta quinque, glabra, post anthesin 1,5 mm longa. Anthera quinque,
1 mm longa, 0,7 mm lata, aurantia. Pollen 52 longum, 30 p latum, oblongum.
Discus ovarium excedens, viridus, nectarifer. Stigma 0,6 mm diam., globosum,
capitatum. Stylus 1,3 mm longus, glaber, erectus. Ovarium 5-carpellatum, 0,7 mm
longum, 0,6 mm diam., apicibus obtusis, lateribus sparsim setulosis. Fructus
5-carpellatus, 6,5 mm longus, 4 mm diam., viridus; cornibus 1 mm longis, erectis,
echinulatis. Semen 3,7 mm longum, arrilatum, 1,6 mm latum, piceum, nitens.
Type: CAPE—3420 (Bredasdorp): north side of pass from Wydgelegen to De Hoop, at
base of limestone hills, on gently sloping ground in crevices and in shallow soil on
limestone bedrock (-AD), 100 m (350 ft.), 3/8/1975, Williams 2028 (NBG, holotype; K,
MO, NSW, PERTH, PRE, S, STE, isotypes).
Euchaetis intonsa was apparently first collected by J. P. H. Acocks in
December 1962.
Shrubs up to 400 mm tall forming a dense rounded bush if undisturbed, single
stemmed at base. Branches fairly numerous, somewhat erect, smooth, glabrescent
as the outer skin splits off, ashy grey. Branchlets numerous, erect, puberulous,
reddened where exposed, fairly slender, well clothed with leaves, irregularly
alternate. Leaves 4 mm long, 2,2—2,4 mm broad, smaller on branchlets, elliptic,
sub-acute, sub-erect, alternate, sometimes 4-ranked on small shoots near the base
of the plant; midrib prominent, gland-dotted, apex callused; abaxial surface
sparsely pubescent mainly on the midrib; margins thick, sparsely ciliate, minutely
gland-dotted; adaxial surface glabrous; petiole 0,5 mm long, adpressed, glabrous,
pale. Inflorescence terminal, 3—4 nate, sessile; florets crowded, flowering simul-
taneously. Bract one to each flower, 2,7 mm long, 1,1—1,7 mm broad, oblanceo-
late, obtuse; apex thick, green, somewhat elevated; margins ciliate, translucent
below; midrib prominent; abaxial surface sparsely pubescent, without gland dots.
Bracteoles two, 2,5 mm long, 1 mm broad, oblong, obtuse; apex thick, green;
margins ciliate; abaxial surface pubescent. Calyx lobes five, 2,6 mm long, 2 mm
broad, oblong, asymmetrical in outline, conspicuous, purplish; apex elevated,
obtuse; margins ciliate, broadly translucent, pinkish; abaxial surface thinly pubes-
cent. Petals five, 3,83—4,2 mm long; limb 1,3-1,8 mm broad irregularly orbicular,
pinkish, becoming recurved; claw 0,8—1 mm broad, strongly transversely bearded,
ciliate above. Staminodes five, 0,1 mm long, vestigial, pale, minutely stalked,
arising on the outside base of the disc. Filaments five, becoming 1,5 mm long,
acicular, glabrous. Anthers five, 1 mm long, 0,7 mm broad, reddish-orange in
colour with a minute spherical apical gland pointed inwards; thecae pointed at
apex and base. Pollen 52 long, 30 ws broad, oblong. Disc exceeds the ovary by
a long way, level on top, bright green, exudes nectar. Stigma 0,6 mm diam.,
globose, capitate. Style becoming 1,3 mm long, glabrous, erect. Ovary
5-carpellate, 0,7 mm long, 0,6 mm diam., apices hemispherically obtuse with a
Studies in genera of Diosmeae (Rutaceae): 7 3377
few hairs at the sides. Fruit (from Williams 2238) 5-carpellate, 6,5 mm long, 4
mm diam., green, glabrous except for a few hairs on the margins and upper
adaxial surfaces; horns + 1 mm long, erect, emarginate; apex bristly with very
short hairs; style persisting. Seed 3,7 mm long including the white aril (0,7 mm
long), 1,6 mm broad, black, shining.
SPECIMENS EX AMINED
CAPE—3420 (Bredasdorp): north side of pass from Wydgelegen to De Hoop,
at base of limestone hills (-AD), 100 m, 3/8/1975, Williams 2028 (NBG, K, MO,
NSW, PERTH, PRE, S, STE), 9/4/1974, Williams 1887 (NBG), 9/11/1976,
Williams 2338 (NBG); 2 miles south of Wydgelegen PO, Kalk Coastal Fynbos,
occasional in valley, 400 ft, 17/12/1962, Acocks 23169 (PRE).
DISTRIBUTION AND BIOLOGY
Up to now Euchaetis intonsa has only been found in one locality on the north
slopes of the limestone hills running from Bredasdorp to Potberg in the vicinity of
Wydgelegen. It would therefore certainly appear to be very rare. Collections of
flowering material made in March, when few flowers were found, and in August
when many buds were found, indicate a rather prolonged flowering period. Ripe
fruits were collected in November. The presence of nectar is one indication that
this plant is insect pollinated. Regeneration after fires would appear to be only
from seed as the plant is single stemmed at base and the root shows no sign of
previous burning off.
VARIATION
Upon encountering a species with such a limited distribution one might expect
it to be a hybrid or variant. However in this case, as no probable parent plants
have been found, hybridisation or variation would appear to be ruled out.
DISCUSSION
The strongly bearded petals most decidedly place this plant in the genus
Euchaetis.
E. intonsa is a distinct species having leaves alternate, elliptic, sparsely
pubescent and ciliate, with bracts, bracteoles and calyx lobes pubescent and with
petals small with the midrib glabrous on the inside. It differs from E. burchellii,
E. laevigata and E. scabricosta which have opposite leaves, glabrous bracts and
glabrous calyx lobes. Although E. meridionalis, a more glabrous plant, is found
nearby and may also have 4 or 5-ranked leaves, it differs from E. intonsa which
has smaller, sparsely pubescent, leaves and pubescent bracts, bracteoles and calyx
lobes. The epithet ‘‘intonsa’’ refers to this hairiness.
338 Journal of South African Botany
"
Fic. 3.
Euchaetis intonsa: 1, leaf. 2, bract. 3, bracteole. 4, calyx lobe. 5, petal. 6, anther. 7,
gynoecium with section of disc. 8, pollen. 9, distribution.
Fic. 4.
Euchaetis intonsa showing the transversely bearded petals.
Studies in genera of Diosmeae (Rutaceae): 7 339
Euchaetis avisylvana Williams, sp. nov. propria propter folia glabra longi-
ciliata, sepala lanceolata acuta alba petaloidea, petala acutissima.
Frutex 1 m, ad basim monocaulis. Rami numerosi, recti, graciles, 2—3-
chotomi, glabri. Ramuli erecti, graciles, recti, glabri, foliosi. Folia 11-14 mm
longa, 1,2—1,4 mm lata, lineari-lanceolata, glabra, acuta, mucronata, longi-ciliata,
erecta, sub-imbricata, alterna, glanduloso-punctata breve petiolata. Inflorescentia
terminalis, ad 19 aggregata, alba. Bractea 7 mm longa, 1—2 mm lata, lineari-
lanceolata, acuta, glabra, glanduloso-punctata, ad apice ciliata. Bracteolae duae,
4,2-4,5 mm longae, 1 mm latae, lanceolatae, acutae, glabrae, sparsim glanduloso-
punctatae. Sepala quinque, 5 mm longa, 1,5-1,8 mm lata, lanceolata, acuta,
glabra, eciliata, alba, petaloidea, persistentes. Petala quinque, 5 mm longa; limbus
2,5 mm longus, 1,1 mm latus, lanceolatus, acutissimus, glaber; unguis 2,5 mm
longus, 1,5 mm latus, ovatus, forte transverse-barbatus, ciliatus. Staminodia
quinque, 0,1 mm diam., vestigialia. Fila quinque, post anthesin 1,3 mm longa,
glabra, acicularis. Antherae quinque, 1 mm longae, 0,7 mm latae, vinosae,
apicibus minute glandulosis. Pollen + 43 w longum, + 21 w latum, oblongum.
Discus 5-sinuato-crenulatus, apertus, pallido-viridus, nectarifer, ovarium exce-
dens. Stigma 0,25 mm diam., pallido-viridum, capitellatum. Stylus initio deflexus,
deinde erectus, 1 mm longus, glaber. Ovarium 5-carpellatum, 0,7 mm longum, 0,8
mm diam., glabrum. Fructus 5-carpellatus, 7 mm longus, 6 mm diam., glaber,
laevis, sparsim glanduloso-punctatus, breve pedicellatus; cornibus 1—1,5 mm
longis, erectiusculis. Semen 4,6 mm longum, 2,4 mm latum, piceum, nitens.
Type: CAPE—3320 (Montagu): Grootvadersbos, on a stony south facing shoulder,
Heidelberg Division (-DD), 425 m (1 400 ft.) alt., 29/4/1975, Williams 1997 (NBG,
holotype; BOL, C, GRA, K, M, MO, NSW, PERTH, PRE, S, STE, isotypes).
This comparatively rare plant was first discovered by Miss Elsie Esterhuysen
in September 1944 on the south slopes of the Langebergen near Strawberry Hill.
This is the name of a farm adjacent to the Grootvadersbos Forest Reserve.
Shrubs 1 m tall, branching from near the base into a diffuse bush spreading
about 1 m diam. Branches numerous, slender, fairly straight, di-trichotomous,
glabrous, brown with the pale outer skin splitting off, smooth but for the small leaf
scars. Branchlets erect, slender, straight, glabrous, pale green, well clothed with
leaves. Leaves 11-14 mm long, 1,2—1,4 mm broad, linear-lanceolate, glabrous,
erect, sub-imbricate alternate; apex acute, incurved, mucronate with a few minute
hairs; margins villous-ciliate and narrowly cartilaginous; adaxial surface some-
what concave; abaxial surface with a row of gland dots on either side of the
midrib; narrowed at the base to a short yellowish petiole. Inflorescence terminal,
up to 9 florets crowded together on short branchlets or pedicels, the most advanced
being nearest to the apex of the main axis. Bract one to each flower, 7 mm long,
1,2 mm broad, linear-lanceolate, acute, glabrous, incurved, tipped with three
minute hairs, margins ciliate above, narrowly translucent below, gland-dotted to
340 Journal of South African Botany
either side of the midrib, sub-sessile. Bracteoles two, 4,2—-4,5 mm long, 1 mm
broad, narrowing evenly to the apex, lanceolate, acute, glabrous, tipped with one
or two minute hairs, margins eciliate and narrowly translucent, sparsely gland-
dotted, sessile. Calyx lobes five, 5 mm long, 1,5—1,8 mm broad, lanceolate, acute,
glabrous, eciliate, white, erect, petaloid, persisting, with about 5 indistinct gland
dots. Petals five, 5 mm long; limb about 2,5 mm long, 1,1 mm broad, glabrous,
lanceolate-acute, recurved, sub-apiculate; claw about 2,5 mm long, 1,4 mm broad,
ovate, strongly transversely bearded across the top, margins crisped ciliate,
delapsing soon after anthesis. Staminodes five, 0,1 mm diam., a vestigial hemis-
pherical lump on the perimeter of the disc. Filaments five, becoming 1,3 mm
long, glabrous, acicular. Anthers five, 1 mm long, 0,7 mm broad, wine coloured
with a small globose apical gland. Pollen + 43 ww, + 21 pw broad, oblong. Disc
5-sinuo-crenulate, more or less erect around the ovary exceeding it somewhat, pale
green, exudes nectar. Stigma 0,25 mm diam., capitellate, pale green. Style
deflexed at first, becoming 1 mm long, erect, glabrous. Ovary 5-carpellate, 0,7
mm long, 0,8 mm diam., glabrous with the carpels leaning inward towards the
apex. Fruit (from Williams 2071) 5-carpellate, 7 mm long, 6 mm diam., glabrous,
smooth, indistinctly gland-dotted, pink tinged below; horns very short, 1—1,5 mm
long, fairly erect, purple tipped. Seed 4,6 mm long, 2,4 mm broad, black, shining;
aril much reduced, black.
SPECIMENS EXAMINED
CAPE—3320 (Montagu): Strawberry Hill, lower south slopes of Langebergen
on the steep slope above kloof (-DD), 2 000 ft., 10/9/1944, Esterhuysen 10406
(BOL, K, NBG); Rocky slopes above stream at Naauwkrans on lower slopes of
Langebergen near Heidelberg, 28/3/1948, Esterhuysen 14444 (BOL, K, NBG);
Strawberry Hill vicinity near Heidelberg, 1 800 ft., 21/7/1952, Wurts 245 (NBG);
Grootvadersbos, 6/4/1959, Barker 8954 (NBG); Langeberg, between Lemoens-
hoek and Naauwkrantz, farm Strawberry Hill, 11/1/1957, Stokoe s.n. (NBG);
Grootvadersbos, stony south facing shoulder, Heidelberg Div., 425 m (1 400 ft.),
29/4/1975, Williams 1997 (NBG, BOL, C, GRA, K, M, MO, NSW, PERTH,
PRE, S, STE); Tradouw Pass, south side near the cave, Heidelberg Div. (-DC),
1 000 ft., 10/9/1975, Williams 2068 (NBG); Tradouw Pass, rocky shoulder about
one third of way up, 1 100 ft., 10/9/1975, Williams 2071 (NBG).
DISTRIBUTION AND BIOLOGY
Euchaetis avisylvana has been found only in a few places on the south slopes
of the Langeberg range from Tradouw Pass eastwards for about 17 kilometres at
altitudes of from 300-600 m (1 000-2 000 ft.). It grows on rocky or stony places
in well drained acid soil derived from the Table Mountain Sandstone. It flowers
from March until about September when ripe fruits may be collected. The
reproductive parts of the flower are well protected by the bearded ciliate petals
Studies in genera of Diosmeae (Rutaceae): 7 341
which however delapse long before the fruit has ripened leaving the pale persistent
calyx lobes which look rather like petals. The flowers have a pungent smell rather
like the excreta of otters, exactly similar to that of E. longicornis, but differing
from that of E. elata which smells of cats’ urine. These facts and the presence of
nectar would indicate that the flowers are most probably pollinated by insects.
Being single stemmed at base, without any stout persistent root, regeneration after
fires can only take place from the seed which is ejected by the usual catapult
mechanism when ripe. This is a factor which could make this plant susceptible to
elimination by too frequent burning of the habitat. As it is, it appears to be in a
precarious position as relic populations in the Tradouw Pass are at present being
eliminated by road building and the population at Grootvadersbos is standing
beneath the trees in a pine plantation belonging to the Department of Forestry.
DIsCUSSION
The strongly bearded petals and the vestigial staminodes are the main charac-
ters which place this plant in the genus Euchaetis.
Euchaetis avisylvana is distinct on account of having villous-ciliate leaves
tipped with a few hairs at the apex; lanceolate-acute, white, somewhat petal-like
Fie. 5.
2, bract. 3, bracteole. 4, calyx lobe. 5, petal. 6, anther. 1,
gynoecium. 8, pollen. 9, distribution.
Euchaetis avisylvana: 1, leaf.
342 Journal of South African Botany
calyx lobes and acute petals. It differs from E. elata E. & Z. which has leaves,
bracts, bracteoles and calyx lobes much shorter and ciliolate not villous-ciliate,
and also the blade of the petal elliptic-apiculate not lanceolate-acute. The very
short horns of E. avisylvana at once distinguish it from E. longicornis Williams
which has very sharp pointed leaves; bracts, bracteoles and calyx lobes with
scattered gland dots. The name is a latinisation of grootvadersbos, the locality
where the type material was gathered.
ae Fic. 6.
Euchaetis avisylvana showing the transversely bearded petals and, with petals removed, the |
disc with vestigial staminodes.
Studies in genera of Diosmeae (Rutaceae): 7 343
Fic. 7.
Euchaetis avisylvana habit.
Acmadenia patentifolia Williams, sp. nov. propria propter folia parva crassa
patentia alterna, flores parvos petalis patentibus, fructum magnum cornibus longis.
Frutex 1 m, 2 m diam., densus, ad basim monocaulis. Rami numerosi, erecti,
breves, glabri, dichotomi. Ramuli numerosi, erecti, brevissimi, sub-flexuosi,
glanduloso-punctati. Folia 1,8—2,7 mm longa, 1,3-1,6 mm lata, oblongo-
lanceolata, obtusa, sessilia, crassa, patentia, alterna, prope basin sparsim puberulis,
2-3 glanduloso-punctata. /nflorescentia terminalis, sessilis, 1-2 nata. Bractea 1,5
mm longa, 1—1,2 mm lata, foliis similis. Bracteolae duae, 1,4 mm longae, 1 mm
latae, late-lanceolatae, obtusae, marginibus ad basin translucentibus sparsim
ciliolatis puberulisque, intus minute puberulae. Sepala quinque, 1,5 mm longa,
1—-1,2 mm lata, oblonga, obtusa, ad apicem crassa, marginibus ad basin anguste
hyalinis sparsim ciliolatis puberulisque, intus minute puberula. Petala quinque,
2,6—2,8 mm longa; limbus 2 mm longus, 1,3 mm latus, lanceolatus, obtusiusculus,
glaber, 1-3 glanduloso-punctatus. Staminodia quinque, 0,8 mm longa, ad apices
glandulosa. Filamenta quinque, post anthesin 1,5 mm longa, acicularia, glabra,
erecta. Antherae quinque, ante anthesin 1 mm longae, 0,6 mm latae, apicibus
sessili-glandulosis. Pollen 43 longum, 23 mw latum, ellipticum. Discus viridus,
crassus, nectarifer, obvallatus. Stigma 0,3 mm diam., capitellatum. Stylus 1 mm
longus, erectus, glaber, persistens. Ovarium 5-carpellatum, 0,8 mm long»m, 0,8
344 Journal of South African Botany
mm diam., glabrum, apicibus globosis. Fructus 5-carpellatus, 16 mm longus, 7
mm diam., glaber, viridus, multi-glanduloso-punctatus, coribus 4 mm longis.
Semen 5—5,6 mm longum arilo exclusa, 2—2,3 mm latum, piceum, nitens; arillus
1—1,5 mm longus, albus.
Type: CAPE—3219 (Wuppertal): on track from Krom River to Breekkrantz, South
Cedarberg, Clanwilliam Division (-CB), 914 m (3 000 ft.), 18/9/1975, Williams 2087
(NBG, holotype; K, MO, M, PRE, S, STE, isotypes).
Acmadenia patentifolia was discovered by Miss Elsie Esterhuysen of the
Bolus Herbarium in 1952 at Krom River in the South Cedarberg. She encountered
this plant again in 1961 on the Swart Ruggens in the Ceres Division. No further
collections were made until 1975 when the material upon which this description
was based was gathered near Krom River in the Clanwilliam Division.
Shrubs 1 m tall, spreading to 2 m diam., forming a dense rounded bush with
many branches arising from a single stem at base. Branches numerous, erect,
short, glabrous, irregularly bent, dichotomous; bark rough, ashy brown. Branch-
lets numerous, erect, very short, sub-sinuous, sparsely and minutely puberulous,
reddish, lumpy with oil glands at open spaces, often encrusted with small pieces of
a red resinous exudate at the base of the leaves. Leaves 1,8—2,7 mm long, 1,3—1,6
mm broad, oblong-lanceolate, obtuse, thick, sessile, spreading, alternate; apex
with an immersed callus; margins rounded with the cartilage immersed; midrib
rounded with 1-3 gland dots; the base thinly puberulous clasping the stem.
Inflorescence terminal, sessile, solitary or paired; flower 6 mm diam., with the
white petals spreading to reveal the green calyx lobes and the green disc; calyx
minutely puberulous. Bract 1,5 mm long, 1—1,2 mm broad, leaf-like, somewhat
ciliolate at base. Bracteoles two, 1,4 mm long, 1 mm broad, broadly lanceolate,
obtuse, 1—3 gland-dotted; margins below broadly translucent, minutely and
sparsely ciliolate, thinly puberulous; adaxial surface minutely puberulous. Calyx
lobes five, 1,5 mm long, 1—-1,2 mm broad, oblong, obtuse, thick at the apex,
sparsely gland-dotted, sparsely and minutely puberulous inside and at the sides
below. Petals five, 2,6—2,8 mm long; limb 2 mm long, 1,3 mm broad, lanceolate,
sub-obtuse, glabrous, 1—3 gland-dotted, white, spreading-recurved, margins mi-
nutely denticulate; claw 0,8 mm long, smooth. Staminodes five, 0,8 mm long,
0,25 mm broad at base, transversely flattened below, arising near the base of the
petal, apical gland 0,15 mm diam., globose, pale yellow. Filaments five, becom-
ing 1,5 mm long, acicular, glabrous, erect. Anthers five, before anthesis 1 mm
long, 0,6 mm broad, apical gland small semi-immersed. Pollen 43 pb long, 23 w
broad, elliptic. Disc erect, green, thick, exudes nectar, slightly exceeds the ovary.
Stigma almost 0,3 mm diam., capitellate. Style becoming 1 mm long, erect,
glabrous, persisting. Ovary 5-carpellate, 0,8 mm long, 0,8 mm diam., glabrous,
apices globose. Fruit 5-carpellate, 16 mm long, 7 mm diam., glabrous, green,
multi-gland-dotted; horns 4 mm long, spreading at +45 °, tinged with purple,
Studies in genera of Diosmeae (Rutaceae): 7 345
apex with a small immersed gland. Seed 5—5,6 mm long excluding the aril, 22,3
mm broad, black, shining; ari] 1—-1,5 mm long, white.
SPECIMENS EXAMINED
CAPE—3219 (Wuppertal): rocky ridge, Krom River, South Cedarberg,
Clanwilliam Division (-CB), 3 000-4 000 ft., 4/10/1952, Esterhuysen 20542
(BOL); on track from Krom River to Breekkrantz, South Cedarberg, Clanwilliam
Division, 914 m (3 000 ft.), 18/9/1975, Williams 2087 (NBG, K, MO, M, PRE,
S, STE); on rocky plateau, Stompiesfontein, Swartruggens, Ceres Division (-DC),
4 000 ft., 19/11/1961, Esterhuysen 29351 (BOL).
DISTRIBUTION, BIOLOGY AND VARIATION
Acmadenia patentifolia, with its insignificant flowers, has very seldom been
gathered. Although dominant in the small areas where it has been found it appears
to be a very rare plant. Being in some respects (i.e. the fruit), remarkably like a
Diosma it would be a pity if this possible link with the Acmadenia were to be lost.
It has been found growing on a stony quartzitic ground derived from Cape
Geological System in, as far as is known, only two localities, about 40 km apart,
both falling within the winter rainfall area of the South Western Cape. No
variation has been noted.
The little white flowers, with spreading petals exposing the nectariferous disc,
are most probably pollinated by insects. The seeds when ripe are ejected by the
usual catapult mechanism and regeneration appears to be only from seed. A
section, 13 mm diam., from the stem near the base of one rather small plant
showed 28 annual rings indicating that fires seldom occur. The leaves have a faint
smell of eucalyptus (blue gum) when crushed.
DISCUSSION
Although the wide open flower and the large fruit with long horns give A.
patentifolia the appearance of a Diosma, the presence of staminodes attached at
the outer base of the disc adjacent to the petals, the shape of the disc itself, closely
surrounding the ovary not open and crenulate, and the rather close resemblance of
the whole plant to Acmadenia tetracarpellata lead one to place it in the genus
Acmadenia. Other species in Acmadenia with similar open flowers and small
staminodes are A. matroosbergensis and A. teretifolia. Factors excluding this
plant from other genera of the Diosmeae are: (1) petals without any transverse
beard; (2) staminodes present; (3) anther with a small apical gland; (4) stigma
capitellate; (5) style short; (6) filament and style glabrous.
It is interesting to note the way in which the base of the leaf is closely
adpressed to the branch around its point of attachment, reminding one of a
parasitic scale insect. Acmadenia patentifolia is recognised as distinct on account
of having very small, thick, alternate leaves, small open flowers and a large fruit
with long horns. It looks rather like A. tetracarpellata which however has
somewhat thinner, opposite leaves and a four-carpellate fruit.
346
Journal of South African Botany
Fic. 8.
Acmadenia patentifolia: 1, leaf. 2, leaf. 3, bract. 4, bracteole. 5, staminode. 6, calyx lobe.
7, petal. 8, anther. 9, gynoecium with section of disc. 10, pollen. 11, distribution.
Fic. 9.
Acmadenia patentifolia showing disc, staminodes etc.
—
Se
Studies in genera of Diosmeae (Rutaceae): 7 347
Acmadenia kiwanensis Williams, sp. nov. propria propter folia alterna lan-
ceolata acuta puberula apicibus minute echinulatis, bracteolas nullimodo
apiculatas, fructum aliquantum pubescentem sepala magnopere excedentem.
Frutex 300-500 mm, ad basin monocaulis. Rami patentes, subnumerosi,
glabri. Ramuli aliquanto numerosi, erecti, albopuberuli. Folia 7-14 mm longa,
2,2—1,5 mm lata, lanceolata vel lineari-lanceolata, acuta, petiolata, glanduloso-
punctata, puberula, alterna, patentes, apicibus minute echinulatis, marginibus initio
ciliolatis. Inflorescentia solitaria, terminalis, 10 mm diam., roseus. Bractea 4 mm
longa, 1,5 mm lata, oblonga, acuta, glabra, ad basin translucens. Bracteolae duae,
3,5 mm longae, 2 mm latae, lanceolatae, acutae, ciliatae, glabrae, ad basin
translucentes. Sepala quinque, 3 mm longa, 2 mm lata, ovato-lanceolata, acuta,
glabra, villoso-ciliata, intus in medio pubescentia. Petala quinque, in toto 6 mm
longa; limbus 2,5 mm lata, obovatus, apiculatus, glaber, roseus, recurvus; unguis
3 mm longus, 1,2 mm latus, pubescens. Staminodia quinque, 1 mm longa, erecta,
glabra, ad apice glandulosa. Filamenta quinque, post anthesin 2,5 mm longa,
acicularia. Antherae quinque, 1,5 mm longae, 0,8 mm latae, flavovirentia,
apicibus glandulosis. Pollen 40 w longum, 33 w latum. Discus erectus, breviter
ovarium excedens, viridus. Stigma 0,7 mm diam., viridum, capitatum. Stylus 2
mm longus, erectus, glaber. Ovarium 5-carpellatum, 0,8 mm diam., sparsim
pubescens. Fructus 5-carpellatus, 6,5 mm longus, 4,5 mm diam., aliquantus
pubescens, sepala magnopere excedens, cornibus c. 1,5 mm longis. Semen 4,5
mm longum, 1,9 mm latum, piceum, nitens.
Type: CAPE—3327 (Peddie): on right hand side of road to Kiwane, 2,6 km from main
Peddie/East London road (-BA), 122 m (400 ft.) alt., 9/3/1976, Williams 2175 (NBG,
holotype; BOL, GRA, K, MO, NSW, PRE, S, isotypes).
Acmadenia kiwanensis was apparently collected for the first time by an
Agricultural Officer in the Native Affairs Department in 1952 who noted that it
was ‘‘prevalent between Keiskamma and Chalumna Rivers within five miles of the
coast’’. Comins found it again in 1956 on ‘‘south west facing slopes above Kiwani
River’. Samples of their material, preserved at both Pretoria and Grahamstown,
looked rather different from any other known Acmadenia and the locality was so
much further to the east than that of any other member of the genus, that it became
imperative for this plant to be further investigated by the collection of fresh
material. The collection of fresh material on a field trip of some 2 260 km resulted
in the publication of this description.
Shrubs 300-500 mm tall, single stemmed at base. Branches fairly numerous,
almost sprawling (perhaps due to trampling by cattle), greyish-brown, glabrous.
Branchlets fairly numerous, erect, somewhat clustered, not hidden by the leaves,
at first densely clothed with short white erect hairs. Leaves 7-14 mm long
including the petiole 1—1,5 mm long, 2,2-1,5 mm broad, the longest being
narrower, lanceolate to linear-lanceolate, acute, 5-ranked, alternate, ©preading,
348 Journal of South African Botany
uppermost erect at first, apex pale minutely bristly; margins scabrid, at first
ciliolate becoming virtually eciliate; abaxial surface with gland dots small and
numerous scattered between midrib and margins, pubescent with short erect hairs
mainly towards the margins and midrib. Inflorescence solitary, terminal, 10 mm
diam., pink, subtended by several bract-like leaves. Bract 4 mm long, 1,5 mm
broad, oblong, acute, glabrous or puberulous, petiolate. Bracteoles two, 3,5 mm
long, 2 mm broad, lanceolate, acute, puberulous above, ciliate, broadly translucent
below. Calyx lobes five, 3,5 mm long, 2 mm broad, ovate-lanceolate, puberulous
above; apex acute green, elevated; margins ciliate, broadly translucent and villous
ciliate below; adaxial surface pubescent in the middle. Petals five, 6 mm long
overall; Jimb 2,5 mm broad, obovate, apiculate, glabrous, pink, spreading; claw 3
mm long, 1,2 mm broad, narrowing to the base, pubescent and bulged inwards at
the throat. Staminodes five, 1 mm long, erect, glabrous, gland-tipped. Filaments
five, becoming 2,5 mm long erect, glabrous, acicular. Anthers five, 1,5 mm long,
0,8 mm broad, greenish-yellow, apical gland pointed and minute. Pollen 40 w
long, 33 w broad, broadly elliptic. Disc fleshy, green, erect, just exceeding the
ovary. Stigma 0,7 mm diam., green, capitate. Style becoming 2 mm long,
glabrous, ever-erect. Ovary 5-carpellate, 0,8 mm diam., somewhat pubescent at
the sides of the carpels. Fruit 5-carpellate, 6,5 mm long, 4,5 mm diam.; carpels
green, minutely gland-dotted, short-pubescent mainly at the sides; horns 1,5 mm
long, erect, apex cleft adaxially. Seed 4,5 mm long, 1,9 mm diam., black, shining,
endosperm black-covered.
SPECIMENS EXAMINED
CAPE—3327 (Peddie): prevalent between Keiskamma and Chalumna rivers,
within 5 miles of coast (-BA), -/8/1952, C.A.O. Native Affairs Dept. A1906
(GRA, PRE); S.W. facing slope above Kiwane river 18/5/1956, Comins 1522
(GRA, PRE); on right hand side of road to Kiwane, 2,6 km from the main
Peddie/East London road 122 m (400 ft.), 9/3/1976, Williams 2175 (NBG, BOL,
GRA, K, MO, NSW, PRE, S).
DISTRIBUTION AND BIOLOGY
Acmadenia kiwanensis, so far as is known, occurs only between the Keis-
kamma and Chalumna rivers within 8 km of the coast. It grows in relic fynbos
surrounded by mixed grassveld in shallow soil on flat quartzitic sandstone. The
surrounding area is fairly heavily grazed by cattle and frequently burnt off. The
plants appear to have been trodden down at some time but have lately escaped
burning. It is doubtful if they could survive veld fires as there is no evidence of a
persistent rootstock such as is possessed by Agathosma peglerae, a plant which
occurs nearby in the recently burnt grassveld. One or two ripe fruits having been
seen early in March, indicate that flowering had been taking place some months
earlier and material collected some time in August was also still flowering. From
Studies in genera of Diosmeae (Rutaceae): 7 349
this it would appear that flowering must extend over a considerable part of the
year. The pink flowers and nectariferous disc indicate that this plant is most
probably pollinated by insects. Regeneration is from seed which is ejected when
ripe in the usual way by a catapult mechanism. The leaves, when crushed, have a
pleasant sweet smell faintly approaching that of aniseed.
DISCUSSION
This plant is placed in the genus Acmadenia because of having clawed petals,
pubescent and bulging inwards at the throat, staminodes 1 mm long and an
obvallate disc, i.e. that exceeds and surrounds the ovary but does not spread
outwards or close over it.
Characters that exclude other genera of the Diosmeae are: (1) style and
filaments glabrous and short; (2) anthers with a small apical gland; (3) fruit
5-carpellate.
26°30
© PEDDIE ss)
ee aT CHALUMNA RIVER
KEISKAMMA RIVER
@ GRAHAMS TOWN
°
33 30
ALFRED
Fic. 10.
Acmadenia kiwanensis: 1, leaf. 2, bract. 3, bracteole. 4, ca
gynoecium with section of disc. 8, pollen.
lyx lobe. 5, petal. 6, anther. 7,
9, distribution.
350 Journal of South African Botany
Acmadenia kiwanensis is recognised as distinct on account of its having
alternate, lanceolate-acute, puberulous leaves with the tips pale and minutely
bristly, fruits somewhat pubescent with fairly short horns and imbricated only
towards the base by the calyx lobes. Geographically the nearest species in the
same genus is A. obtusata (Thunb.) B. & W., about 200 km to the south west,
near to Port Elizabeth, growing on limestone, with leaves nearly always opposite,
with bracts and bracteoles apiculate and with fruits glabrous, short horned and
more or less hidden by the calyx lobes. Further to the west beginning near
Plettenberg Bay A. alternifolia can be readily distinguished by its fruits with much
longer horns.
Acmadenia nivea Williams, sp. nov. propria propter folia lanceolata glabra,
petala nivea, staminodia 0,09—0,5 mm variantia, antheras minute + sessili-
glanduliferas, ovarium 3 vel 4-carpeilatum, cornibus binatim perbrevibus.
Frutex 200-300 mm, graciles, diffusus, ad basin monocaulis. Rami pauci,
laxiusculi, glabrescenti, porphyrei, sparsim 2-chotomi. Ramuli pauci, graciles,
erecti, sparsim puberuli, porphyrei. Folia 6-8 mm longa, 1,5—1,8 mm lata,
lanceolata, acuta, rubro-mucronata, brevi petiolata, glabra, bifaria glanduloso-
punctata, adpressa vel sub-patentia, alterna vel interdum ad basin opposita,
marginibus late translucentibus ciliatis vel eciliatis. Inflorescentia terminalis, ad
7-aggregata; flos 6 mm diam., sessilis, niveus, fauce contractus. Bractea 2,7—5
mm longa, 0,7/—1,3 mm lata, extima folio simili, intima lanceolata, rubro-
mucronata, glabra, glanduloso-punctata, sessilia, marginibus ad basin late trans-
lucentibus ciliolatis. Bracteolae duae, 2,3 mm longae, 0,6 mm latae, lanceolatae,
rubro-mucronatae, ciliolatae, marginibus ad basin translucentibus. Sepala quinque,
2,8-3 mm longa, 1,5 mm lata, elliptica, glabra, ciliata, rubro-mucronata, mar-
ginibus late translucentibus. Petala quinque, 5 mm longa; limbus 1,8 mm latus,
ellipticus, glaber, nivea, patens; unguis glaber, translucens, ciliolatus vel
eciliolatus. Staminodia quinque, 0,09-0,5 mm longa variantia, vestigialia.
Filamenta quinque, post anthesin 1 mm longa, glabra, acicularia, erecta. Antherae
quinqgue, ante anthesin 0,9 mm longae, 0,6 mm latae, vinosae, minute glan-
duliferae. Stigma 0,3 mm diam., viridum, capitatum. Stylus 0,5 mm longus,
glaber, erectus. Ovarium 3 vel 4-carpellatum, 0,5 mm diam., globosum, glabrum;
carpellum ad apicem bifidum. Fructus 3 vel 4-carpellatus, breve pedicellatus, 3,5
mm longus, glaber; cornibus binatim, perbrevibus. Semen 2,3 mm longum, 1,5
mm latum, ovoideum, piceum, nitens.
Type: CAPE—3418 (Simonstown): marshy ground, peaty soil, amongst restiads,
Steenbras catchment area, Kogelberg, Caledon Division (-BB), 1 265 m (4 150 ft.),
13/6/1975, Williams 2006 (NBG, holotype).
Acmadenia nivea was first discovered by T. Stokoe in August 1924 on the
Kogelberg. He also found it on the top of Somerset Sneeuwkop in 1937. Being a
Studies in genera of Diosmeae (Rutaceae): 7 351
rare plant it has been very seldom seen and very little material has been
distributed.
Shrubs 200-300 mm tall, slender, diffuse, branching from a single stem at
base. Branches few, somewhat lax, glabrescent, reddish-brown, sparingly
dichotomous. Branchlets few, slender, erect, sparsely puberulous, reddish-brown.
Leaves 6-8 mm long, 1,5—1,8 mm broad, lanceolate, acute, rubro-mucronate,
short-petiolate, glabrous, with 2 rows of gland dots, adpressed or sub-spreading,
alternate or at the base sometimes opposite; margins broadly translucent, thin,
ciliate or eciliate. Inflorescence terminal, up to 7-aggregate; flower 6 mm diam.,
sessile, pure white, contracted at the throat. Bract 2,7—-5 mm long, 0,7—1,3 mm
broad, the outermost leaf-like, the inner becomingly lanceolate, rubro-mucronate,
glabrous, gland-dotted, sessile; margins towards the base broadly translucent,
ciliolate. Bracteoles two, 2,3 mm long, 0,6 mm broad, lanceolate, rubromucro-
nate, ciliolate; margins translucent below. Sepals five, 2,8-3 mm long, 1,5 mm
broad, elliptic, glabrous, ciliate, rubro-mucronate; margins broadly translucent.
Petals five, 5 mm long bent inwards at the throat; /imb 1,8 mm broad, elliptic,
glabrous, pure white, spreading; claw glabrous, translucent, ciliate above or
eciliate. Staminodes five, varying from 0,09 mm to 0,5 mm vestigial, arising on
the outside of the disc. Filaments five, becoming 1 mm long, glabrous, acicular,
erect. Anthers five, 0,9 mm long, 0,6 mm broad before anthesis, wine coloured
with a minute apical gland. Stigma 0,3 mm diam., depressed globose, green,
capitate. Style becoming 0,5 mm long, glabrous, erect. Ovary 3-carpellate or (as
in Esterhuysen 15245) 4-carpellate, 0,5 mm diam., globose, glabrous; carpel bifid
at the apex. Fruit 3—4 carpellate, short-pedicellate, 3,5 mm long, glabrous; horns
paired, extremely short. Seed 2,3 mm long, 1,5 mm broad, egg-shaped, black,
shining.
SPECIMENS EXAMINED
CAPE—3418 (Simonstown): Kogelberg, near the summit, marshy areas,
Caledon Division (-BB), 4 100 ft., —/8/1924, Stokoe 968 (BOL), —/11/1944,
Stokoe s.n. (BM, BOL, NBG, PRE, SAM 58809), 15/11/1964, Esterhuysen 30644
(BOL), 7/5/1966, Esterhuysen s.n. (BOL), 15/4/1966, Thompson 181 (PRE,
STE), 21/4/1970, Boucher 1244 (STE), 13/6/1975, Williams 2006 (NBG); marshy
areas between Kogelberg and Spinnekopnes, 31/9/1975, Vogelpoel s.n. (Williams
2090) (NBG); Somerset West Sneeuwkop, Caledon Division, 5 000 ft., 12/3/1937,
Stokoe 5016 (BOL, SAM 52244), 10/2/1957, Esterhuysen 20764 (BOL); on rock
or on steep rocky slopes on south side of Somerset West Triplets, Caledon
Division, 4 800 ft., 10/4/1949, Esterhuysen 15245 (BOL); Kogelberg Forest
Reserve, north east slopes of Voorberg, Caledon Division (-BD), 550 m,
2/10/1971, Boucher 1648 (STE).
352 Journal of South African Botany
DISTRIBUTION, BIOLOGY AND VARIATION
Acmadenia nivea is found only on the high peaks of the Hottentots Holland
mountains 1 250 to 1 600 m (4 100—5 200 ft.) above sea level growing in marshy
areas usually facing south to south east. It is an extremely rare plant, a few
individuals only being found scattered amongst the vegetation. It is destroyed by
fires and regeneration can only take place from seed. The brilliantly white petals
must be attractive to insects and perhaps moths. The disc is fairly flat on top, dark
green, fleshy and exudes nectar. It forms an annulus around and slightly above the
unextended stigma while the pollen is being shed. The anthers remain below the
contracted throat of the perianth. After the pollen is shed the style lengthens
placing the receptive stigma on a level with the empty anthers. It would appear
therefore that this plant is insect pollinated and that the reproductive parts of the
flowers are well protected against the inclement weather usually experienced at
these exposed altitudes.
There seems to be some evidence of variation in this species perhaps
indicating a trend towards reduction. Populations from the Somerset Sneeuwkop
area have flowers with staminodes 0,4-0,3 mm long and ovaries with 4 to 3
carpels. Populations from the Kogelberg area have flowers with staminodes
reduced to between 0,3 and 0,1 mm long and ovaries with 3 carpels. In the very
closely related species, A. candida, the staminodes are reduced to a minute scale
0,05 mm in diameter and the ovary to only two carpels.
DISCUSSION
Although having an ovary with only 3—4 carpels instead of the more normal
number of 5, this plant is placed in the genus Acmadenia because of having (1)
petals clawed and bent inwards at the throat without any transverse beard; (2)
staminodes present; (3) anthers with a small apical gland; (4) disc closing over
the ovary; (5) stigma capitellate; (6) style short, erect, glabrous; (7) filaments
short, glabrous.
Acmadenia nivea is recognised as distinct on account of having glabrous,
lanceolate leaves, snow white petals, staminodes varying from 0,09 to 0,5 mm
long, anthers with minute + sessile apical glands and a 3—4 carpellate ovary. It is
nearest to Acmadenia candida, a smaller plant, in which however the ovary is
reduced to only two carpels. The name refers to the pure white petals.
Acmadenia candida Williams, sp. nov. propria propter fruticem gracillimum
glaberrimumque, petala candida, staminodia mimima, antheras glanduliferas mi-
nute stipitatas, ovarium 2-carpellatum.
Frutex 250-300 mm, gracillimus, ad basin monocaulis. Rami pauci, gracil-
limi, laevi, erecti, aphylli, porphyrei, 1-2 chotomi. Ramuli pauci, filiformi, brevi,
sparsim foliosi. Folia 4-5 mm longa, 1 mm lata, lanceolata, glabra, sessilia,
erecta, alterna, minute scabrido-mucronulata, bifaria glanduloso-punctata, sparsim
Studies in genera of Diosmeae (Rutaceae): 7 353}
Fic. 11.
Acmadenia nivea: 1, leaf. 2, bract. 3, bracteole. 4, calyx lobe. 5, petal. 6, anther. 7.
gynoecium. 8, pollen. 9, distribution.
ciliolata vel eciliata, marginibus late translucentibus. Inflorescentia terminalis, +
6-aggregata; flos 4 mm diam., candidus, fauce contractus. Bractea 1,7—2,3 mm
longa, 0,6—0,8 mm lata, lanceolata, acuta, mucronulata, glanduloso-punctata,
glabra, sparsim ciliolata, marginibus translucentibus. Bracteolae duae, 1,1 mm
longae, 0,4 mm latae, lanceolatae, mucronulatae, glabrae, sparsim ciliolatae,
marginibus translucentibus. Sepala quinque, 1,8 mm longa, 0,9 mm lata, ovato-
lanceolata, mucronulata, glabra, sparsim glanduloso-punctata, ciliolata, marginibus
late translucentibus. Petala quinque, 3,1 mm longa; limbus 2 mm diam., or-
bicularis, glaber, candidus, patens; wnguis 1,1 mm longus, 0,7 mm latus, crispo-
ciliatus. Staminodia quinque, 0,05 mm diam., ultima vestigialia. Filamenta
quinque, post anthesin 0,6 mm longa, glabra, erecta. Antherae quinque, ante
anthesin 0,5 mm longae, 0,3 mm latae, vinosae, glanduliferae minute stipitatae.
Pollen 28 u longum, 17 pu latum, oblongum. Stigma 0,15 mm diam., capitellatum,
globosum. Stylus 0,6 mm longus, glaber, erectus. Ovarium 2-carpellatum, + 0,3
mm diam., glabrum; carpellum ad apicem bifidum. Fructus et semen a nobis non
visa.
Type: CAPE—3419 (Caledon): in a marsh on level area, top of road from Nuweberg
Forest Station, mixed up with low dense Restionaceae, east slopes of Landdrost Kop,
Caledon Division (-AA), 1 070-1 220 m (3 500-4 000 ft), 17/3/1968, Esterhuysen 31946
(BOL, holotype).
354 Journal of South African Botany
Acmadenia candida was discovered by T. Stokoe in 1943 growing on the east
slopes of Landdrost Kop in the Hottentots Holland Mountains. It was not until
1968, after the Forestry Department had constructed a road up to the east slopes of
Landdrost Kop, that Miss Elsie Esterhuysen again found this plant. Two months
later she made another collection and subsequently the Forestry Department burnt
the whole area off as a firebreak. Repeated searches in recent years have failed to
reveal anything and now the Forestry Department, having declared this a “*Wil-
derness Area’’, have constructed an enormous mountain chalet at the end of their
access road. It is quite possible that this species is now extinct.
Shrubs 250-300 mm tall, slender, single-stemmed below. Branches few,
extremely slender, smooth, erect, leafless, reddish-brown, 1—2 chotomous. Branch-
lets few, filiform, short, rather sparsely leafy, exoderm like snake skin splitting off,
arising from immediately below the inflorescence. Leaves 4—5 mm long, | mm
broad, lanceolate, scabrido-mucronulate, glabrous, 2-fariously gland-dotted, very
sparsely ciliolate or eciliate, adpressed erect, alternate margins broadly translu-
cent. Inflorescence terminal, up to 6-aggregate with flowers about 4 mm diam.,
pure white, contracted at the throat. Bract 1,7—2,3 mm long, 0,6—0,8 mm broad,
lanceolate, acute, mucronulate, gland-dotted, glabrous, sparsely ciliolate, margins
translucent. Bracteoles two, 1,1 mm long, 0,4 mm broad, lanceolate, mucronulate,
glabrous, sparsely ciliolate, margins translucent. Calyx lobes five, 1,8 mm long,
0,9 mm broad, ovate-lanceolate, mucronulate, glabrous, gland-dotted, ciliolate,
margins broadly translucent. Petals five, 3,1 mm long, clawed, bent inwards at the
throat; /imb 2 mm diam., orbicular, glabrous, spreading, pure white; claw 1,1 mm
long, 0,7 mm broad, margins above shaggy ciliate. Staminodes five, 0,05 mm
diam., an absolutely minute vestigial scale on the outside of the disc. Filaments
five, becoming 0,6 mm long after anthesis, glabrous, erect. Anthers five, 0,5 mm
long, 0,3 mm broad before anthesis, reddened with a minutely stipitate apical
gland. Pollen 28 w long, 17 uw broad, oblong with a rather thick skin. Disc closes
over the ovary to some extent. Stigma 0,15 mm diam., globose, capitellate. Style
becoming 0,6 mm long, glabrous, erect. Ovary 2-carpellate, + 0,3 mm diam.;
carpels glabrous, bifid at the apex. Fruit and Seed not seen.
SPECIMENS EXAMINED
CAPE—3419 (Caledon): east of Landdrost Kop (-AA), -/4/1934, Stokoe 2845
(BOL); in a marsh on level area, top of road from Nuweberg Forest Station, east
slopes of Landdrost Kop 3 500-4 O00ft, 17/3/1968, Esterhuysen 31964 (BOL,
holotype); in a marsh at east base of Landdrost Kop, Nuweberg Forest Reserve
3 000-3 500 ft, 18/5/1968, Esterhuysen 31961 (BOL).
DISTRIBUTION, BIOLOGY AND VARIATION
Acmadenia candida is apparently a very rare plant. As far as is known it
occurred in only one locality on the eastern slopes of the Hottentots Holland
Studies in genera of Diosmeae (Rutaceae): 7 355
mountains at an elevation of about 1 070 m (3 500 ft) above sea level growing
amongst low Restionaceae in a marsh. Flowering appears to extend from mid
March to the middle of May. Fruiting material has not been seen. The conspicuous
pure white flower with a constricted throat, may indicate that it is insect
pollinated, perhaps by some small moth.
With so little material it is impossible to observe any variation but in this
species we see an example of reduction carried to limits that may lead to
extinction.
DISCUSSION
The characters that indicate that this plant should be placed in the genus
Acmadenia and which exclude it from other genera of the Diosmeae are: (1)
petals clawed, bent inwards at the throat without any transverse beard; (2)
staminodes present on the outer surface of the disc although only vestigial; (3)
anthers with a small apical gland; (4) disc closing over the ovary to some extent;
(5) stigma capitellate; (6) style short, erect, glabrous; filaments short, glabrous.
This species is unusual in that it shows almost complete reduction of its
staminodes, in that the apical gland on the anther is minutely stipitate and in that
the ovary is reduced to 2 carpels. These atypical characters do not really serve to
place this species into either Diosma with no staminodes, Adenandra with
stipitate anther glands or even into Agathosma which contains some 2-carpellate
species.
mm
wt >
2,5mm
5mm
oO
2,5mm
50u
Fic. 12.
Acmadenia candida: 1, leaf. 2, bract. 3, bracteole. 4, calyx lobe. Sb petal. 6, anther. 7,
gynoecium with section of disc. 8, pollen. 9, distribution.
356 Journal of South African Botany
Acmadenia candida is closest to Acmadenia nivea which however has
staminodes somewhat less reduced and an ovary with 3 or 4 carpels.
Acmadenia candida is recognised as distinct on account of being an extremely
slender glabrous plant with few branches, with white petals, with reduced
staminodes, with anthers minutely stipitate-glandular and with a 2-carpellate
ovary. The name alludes to the strikingly white petals.
Macrostylis ramulosa Williams, sp. nov. propria propter ramos ad basin
ramulosos, folia lineari-lanceolata pubescentia ecostata, bracteas 2,5—4,5 mm
longas pubescentes, bracteoles et sepalia ad apices pubescentes, fructum pubescen-
tem.
Frutex inferior, 300 mm, sub-decumbens, ad basin monocaulis. Rami sub-
decumbentes, glabrescentes, ad basin ramulosi. Ramuli numerosi erectiusculi,
foliosi, sparsim pubescentes. Folia 6-8 mm longa, 1,2 mm lata, lineari-lanceolata,
acuta, erecta, sessiles, alterna, pubescentia, ecostata, indistincte glanduloso-
punctata. Inflorescentia terminalis, 5-6 aggregata; flos 5 mm diam., albus.
Bractea 2,5—4,5 mm longa, 0,8—1 mm lata, similis folio. Bracteolae duae, 1,6—2
mm longae, 0,6 mm latae, oblongae, sub-acutae, apicibus caespitosis, ad basin
translucentes. Sepala quinque, 3 mm longa, | mm lata, anguste lanceolata, ciliata,
ad basin translucentia, apicibus pubescentibus. Petala quinque, 5 mm longa, alba;
limbus 1,4 mm latus, patens, sub-orbicularis; unguis 1,2 mm latus barbatus,
eciliatus. Staminodia quinque, 0,3 mm longa, saepe vix visibilia. Filamenta
quinque, post anthesin 6—7 mm longa, glabra. Antherae quinque, 1,2 mm longae,
0,7 mm latae, oblongae, flavidae, apicibus minute glandulosis. Pollen 52 w
longum, 22 mw latum, oblongum. Discus ovarium includens, stylus ad basin
cingens, viridus, nectarifer. Stigma simplex, terminale. Stylus 6 mm longus,
glaber. Ovarium 3-carpellatum, 0,6 mm longum, globosum, glabrum. Fructus
(Williams 2261) 3-carpellatus, fere abortu 1-carpellatus, 12 mm longus, 3 mm
latus, pubescens, rubro-glanduloso-punctatus, cornibus erectis, gracilibus. Semen
4,2 mm longum, 1,8 mm latum, arilatum, piceum, impolitum.
Type: CAPE—3219 (Clanwilliam): north facing gravelly slopes, Middelberg Pass near
Kleinplaas, Citrusdal (-CA), 777 m (2 550 ft.), 12/7/1975, Williams 2017 (NBG, holotype;
BOL, K, M, MO, PRE, S, STE, isotypes).
Stokoe collected this plant twice during 1940 and thereafter again in 1945,
1950 and 1955. In spite of its rather distinct hairy appearance it has remained
undescribed until now.
Undershrubs 300 mm tall, semi-decumbent, arising from a single stem almost
below ground level. Branches tending to sprawl, glabrescent, greyish-brown, with
small leaf scars. Branchlets numerous, fairly erect, sparsely pubescent, well
clothed with leaves, many sprout from the older wood. Leaves 6—8 mm long, 1,2
mm broad, linear-lanceolate or subulate, incurved-erect, sessile, alternate, apex
Studies in genera of Diosmeae (Rutaceae): 7 35) 1/
pubescent and acute without any callus, ciliate almost without any hyaline
margins; adaxial surface glabrous, concave; abaxial surface pubescent with scat-
tered long hairs, round backed with many indistinct gland dots scattered mainly
towards the midrib. Inflorescence terminal, 4-6 aggregate; flowers 5 mm diam.,
white. Bract 2,5—4,5 mm long, 0,8—1 mm broad, the inner smaller, similar to the
leaf but with broad translucent margins towards the base. Bracteoles two, 1,6—2
mm long, 0,6 mm broad, oblong, apex sub-acute and tufted with bristly hairs,
ciliate above, margins broad and translucent. Calyx lobes five, 3 mm long, 1 mm
broad, narrowly lanceolate, apex a blunt hairy point, ciliate, margins translucent
below. Petals five, 5 mm long, white; limb 1,4 mm broad, ovate, sub-acute,
spreading; claw 1,2 mm broad, crisped-bearded above in the centre, eciliate.
Staminodes five, 0,3 mm long, slender with a minute apical gland, arising at the
outside base of the disc at the base of the petal, in many cases almost invisible.
Filaments five, glabrous, becoming 6—7 mm long after anthesis. Anthers five, 1,2
mm long, 0,7 mm broad, yellow, oblong with a small apical gland. Pollen 52 w
long, 22 w broad, oblong. Disc green, envelops the ovary, touching the base of
the style, exudes nectar. Stigma simple, terminal. Style becoming 6 mm long,
glabrous, pink at the tip. Ovary 3-carpellate, 0,6 mm diam., globose, glabrous.
Fruit (from Williams 2261) 3-carpellate with normally only one carpel develop-
ing, 12 mm long, 3 mm broad, pubescent, reddish gland-dotted; horns 5 mm long,
erect, slender. Seed 4,2 mm long, 1,8 mm broad, truncate with the aril extending
1,3 mm, black, matt.
SPECIMENS EXAMINED
CAPE—3219 (Clanwilliam): waterfalls between Citrusdal and Elandskloof
(-CA), 3/8/1940, Stokoe 8181 (BOL); west end of Elandskloof above Citrusdal,
5/10/1946, Esterhuysen 13018 (BOL); Elandskloof, -/12/1940, Stokoe s.n. (SAM
58851), 26/3/1951, Compton 22696 (NBG), 27/8/1954, Lewis 4730 (SAM); north
facing gravelly slopes, Middelberg Pass near Kleinplaas, Citrusdal, 777 m (2 550
ft.) 12/7/1975, Williams 2017 (NBG, BOL, K, M, MO, PRE, S, STE); Cedarberg,
Duivelskop, 2 500-3 500 ft. -/1/1945, Stokoe s.n. (BOL, SAM 5881/2);
Duivelskloof, -/9/1950, Stokoe s.n. (SAM 64158).
DISTRIBUTION AND BIOLOGY
Macrostylis ramulosa has been found only in a fairly small area, about 12 km
in length, in the mountains to the east of Citrusdal in the Clanwilliam Division. It
grows on gravelly-sandy soil derived from the Table Mountain series at altitudes
of around 777 m (2 500 ft.) on north facing slopes. Flowers have been seen from
March to September and fruits in December and January.
The disc exudes nectar which rises between the lower parts of the style and the
filaments by capillary action. The small flowers are made more obvious by being
aggregated into groups of 4 to 6. By the time that the anthers shed their pollen
358 Journal of South African Botany
they have been carried far beyond the throat of the flower and likewise, at a later
stage when all the pollen has been shed, the style with the receptive stigma
protrudes a long way. With the above facts in mind it would appear therefore that
this plant is insect pollinated. Regeneration after fires seems to take place only
from seed and not at all by coppicing from the root. The leaves have a very faint
smell of cedar when crushed.
DISCUSSION
This species falls quite naturally into the genus Macrostylis of Bartling and
Wendland. The following characters, besides being typical of the genus, also help
to exclude it from some other genus of the Diosmeae: (1) heads up to 6 aggregate;
(2) petals with a dense shaggy pubescence at the throat; (3) staminodes vestigial,
at most 0,3 mm long; (4) anthers with a minute apical gland; (5) disc closes over
the ovary almost touching the base of the style; (6) stigma simple; (7) style and
filament glabrous, protruding a long way; (8) ovary 3-carpellate.
Macrostylis ramulosa is recognised as distinct on account of being a semi-
decumbent shrub with many branchlets sprouting towards the base, with linear-
lanceolate pubescent leaves without any prominent midrib, with pubescent bracts
2,5—4,5 mm long, with bracteoles and calyx lobes pubescent at their tips, and with
the fruit pubescent. It is nearest to M. decipens which is a much more glabrous
plant with the ability to coppice from the root after fires
lene, JB}.
Macrostylis ramulosa: I, leaf. 2, bract. 3, bracteole. 4, calyx lobe. 5, petal. 6, anther. 7,
gynoecium with section of disc. 8, staminode. 9, pollen. 10, distribution.
Studies in genera of Diosmeae (Rutaceae): 7 359
lI
Fic. 14.
Macrostylis ramulosa showing bearded petals and protruding filaments.
Acmadenia wittebergensis (Compton) Williams, comb. nov.
Diosma _ wittebergensis Compton in Trans.R.Soc.S.Afr. 19.3: 297 (1931).
Type : 3320 (Montagu): Witteberg (Whitehill), south slope, Laingsburg Division
(-BC), 1 067 m (3 500 ft.), 11/8/1924 Compton 2524 (BOL, lectotype).
This plant was discovered by Professor R. H. Compton during the period 1923
to 1929 when he was making collections, in the vicinity of the Karoo Garden at
Whitehill near Matjiesfontein, for his ‘‘Flora of the Whitehill District”. Late in
1974 E. G. H. Oliver found the same plant on the Roodeberg in the Little Karoo
near Ladismith. Examination of this material conveyed the impression that this
species might be more satisfactorily placed in Acmadenia and fresh material was
therefore collected (Williams 2132) to be used as the basis for the following
description.
Shrubs up to 500 mm tall with many stems arising from a persistent rootstock.
360 Journal of South African Botany
Branches erect, stiff, 2—3 chotomous; bark greyish-brown, rough with leaf scars.
Branchlets erect, stiff, short, puberulous, brownish, 2—3 chotomous, leafy. Leaves
11-13 mm long including petiole 1,5 mm long, 1,6-1,9 mm broad, linear-
lanceolate, incurved-erect, alternate, crowded, loosely imbricate; adaxial surface,
lower part of abaxial surface and petiole all puberulous; abaxial surface rounded,
lumpy with two rows of gland dots, scabrid above; apex tapering to a pungent
mucro; base narrowing to a short decurrent petiole; margins rounded, scabrid-
ciliate. Inflorescence terminal, + 5 aggregate in a much reduced raceme of which
several flowers may abort, base somewhat surrounded by the uppermost leaves;
flowers 9 mm diam., white, opening in succession, crowded. Bract one to each
flower, 2,5—4,5 mm long, uppermost shortest, 1 mm broad, lanceolate-subulate,
incurved, with a pungent mucro, erect-pubescent on all sides, round-backed with 2
rows of gland dots. Bracteoles two, 1 mm long, 1,1—1,3 mm broad, deltoid-
orbicular, obtuse, 2—4 gland-dotted, pubescent on both surfaces, margins shaggy-
ciliate. Calyx lobes five, 2,5—2,7 mm long, 1,6—2 mm broad, broadly lanceolate,
acute with a small mucro, pubescent on both surfaces, obscurely gland-dotted,
shaggy-ciliate, reddish. Petals five, 5,8 mm long, 2,7—2,9 mm broad, elliptic,
white, glabrous, recurved, 7 gland-dotted in 2 rows, margins irregularly serrate,
clawed. Staminodes five, 0,5 mm long, conical, spreading-erect, arising close to
the petal on the perimeter of the disc. Filaments five, becoming 1,5 mm long,
glabrous, acicular, incurved. Anthers five, before anthesis 1,3 mm long overall,
0,9 mm broad, very dark red, produced upwards into a thickish support to the
apical gland which produces an exudate before the bud opens. Pollen 40 pw long,
30 w broad, sub-spherical. Disc thick, purplish, minutely pitted, sinuate, almost
level equalling and closely surrounding the ovary, exudes nectar. Stigma 0,3 mm
diam., globose, capitellate, very dark red. Style becoming 1,2 mm long, glabrous,
persisting, slightly deflexed. Ovary 5-carpellate, 1 mm long, 1,2 mm diam.,
glabrous; apices glandular-globose. Fruit 5-carpellate, 8 mm long, 9 mm diam.,
green with purplish tips, multi-gland-dotted; horns extremely short, truncate, apex
emarginate clasping an immersed gland; calyx lobes persisting. Seed 6 mm long,
2,8 mm broad, black, shining, aril large and white.
SPECIMENS EXAMINED
CAPE—3320 (Montagu): Witteberg (Whitehill), south slope Laingsburg Divi-
sion (-BC), 3 500 ft., 11/8/1924, Compton 2524 (BOL); Witteberg, summit,
5 000 ft., 21/5/1925, Compton 2951 (BOL); Witteberg near Whitehall, north
slope, 4 000 ft., 24/10/1943, Compton 15248 (BOL, NBG); Anysberg, rocky
north slopes, 3 000-3 500 ft., 19/4/1941, Stokoe 8187 (BOL, PRE, NBG, SAM
56858), 20/5/1972, Esterhuysen 32860 (PRE); Anysberg, south slopes (-DA),
2 500 ft., 2/8/1956, Wurts 1418 (NBG); Touwsberg, ridge on south slopes (-DB),
3 000-4 000 ft., Esterhuysen 25884 (BOL, NBG); Prinspoortberg, Little Karoo
(-BD), 4 000 ft., -/7/1937, Levyns 6159 (SAM).
Studies in genera of Diosmeae (Rutaceae): 7 361
—3321 (Ladismith): ridge between Rooiberg and Bailey Peak (-CB), 4 500 ft.,
11/11/1974, Oliver 5429 (STE, PRE); Bailey Peak, Rooiberg, north slopes,
4 300/4 800 ft., 2/11/1975, Williams 2132 (NBG).
DISTRIBUTION AND BIOLOGY
Acmadenia wittebergensis has been found at altitudes of from 900—1 500 m
(3 000-5 000 ft.) above sea level growing on dry rocky slopes in quartzitic soil
derived from the Witteberg and Table Mountain series. The distribution is
discontinuous with populations, showing little or no variation, situated on isolated
quartzitic mountains. The surrounding land consisting of softer shales of the
Bokkeveld series has been weathered down to much lower levels. As one cannot
imagine any method of dispersal which could possibly bridge these gaps at the
present time, one is forced to the conclusion that this species, in its present form,
may have been in existence for a very long time. Plants from the Rooiberg seem to
have leaves with long sharp points. The conspicuous white petals in the flower
heads and the presence of nectar indicate that this plant is most probably insect
pollinated. The seeds when ripe are ejected from the capsule by the usual catapult
mechanism but regeneration after fires would appear to take place mainly by
regrowth from a persistent rootstock. One fairly large plant, with about 50 stems,
showed at least four episodes in the branching at the base indicating four past
Fic. 15.
Acmadenia wittebergensis: 1, leaf. 2, bract. 3, bracteole. 4, calyx lobe. 5, petal. 6, anther.
7, staminode. 8, gynoecium with section of disc. 9, pollen. 10, distribution.
362 Journal of South African Botany
destructions by fire. A vigorous young branchlet, springing from the base, may be
quite long, densely clothed with leaves and bear a cluster of twelve flowers at the
apex. The flowers are sweet smelling. The leaves have a resinous smell when
crushed and the fruits a smell of celery when crushed.
This plant has been transferred to the genus Acmadenia because of having its
petals clawed (the claws more or less form a tubular perianth 4 mm long above
which the blades of the petals bend outwards), also because of having pointed
staminodes and a pointed gland at the apex of each anther; the disc does not
spread widely with a crenulate margin as in Diosma but surrounds the ovary rather
closely and is almost level on top. Other factors which exclude Acmadenia
wittebergensis from other genera of the Diosmeae are: (1) petals without any
beard; (2) stigma capitellate; (3) style and filaments short and glabrous; (4) ovary
5-carpellate.
Acmadenia wittebergensis is recognised as being distinct because of having
thick pointed crowded leaves, pubescent bracts, bracteoles and calyx lobes, short
conical staminodes with conical glands on the anthers and a multi-stemmed habit.
It appears to have no very close resemblance to any other species of Acmadenia.
At best one may say that it is nearest to A. teretifolia which Phillips also
transferred to Diosma.
JIS. Afr. Bot. 44 (4): 363-371 (1978)
SS ee ee
HABITAT OF THREATENED SPECIES OF SERRURIA AND PROTEA
ENDEMIC TO WESTERN CAPE COASTAL FLATS
A. V. MILEWSKI*
(Rare and Endangered Plant Species Research Group, Bolus Herbarium, Univer-
sity of Cape Town)
ABSTRACT
The habitats of four species of Serruria and two species of Protea, all threatened
endemics in Coastal Fynbos and Coastal Rhenosterbosveld, are described. These habitats
are compared with those of other threatened species of Proteaceae and Restionaceae.
UITTREKSEL
HABITAT VAN BEDREIGDE SERRURIA EN PROTEA SPESIES ENDEMIES AAN DIE
WESTELIKE KAAPSE KUSVLAKTES
Die habitatte van vier Serruria spesies en twee Protea spesies word beskryf en vergelyk
met die habitatte van ander Proteaceae en Restionaceae spesies. Hierdie bedreigde spesies
kom met beperkte verspreidings in kusfynbos en kusrenosterbosveld voor.
INTRODUCTION
Several recent papers have described the habitats of rare and threatened species
of Proteaceae and Restionaceae on the western Cape coastal flats (Milewski &
Esterhuysen, 1977; Milewski, 1977, Milewski, 1978). This paper describes the
habitats of six additional threatened species of Proteaceae, in relation to those of
the species previously dealt with in this way. Knowledge of the habitats of
threatened species is an urgent prerequisite for their conservation.
METHODS
Six species were chosen for study, each restricted to a small geographical
range of less than 700 km? on the western coastal flats of the south-western Cape
(Milewski & Esterhuysen, 1977; Milewski, 1978). The study species were
Serruria trilopha Knight, S. linearis Salisb. ex Knight, S. furcellata R. Br., S.
cyanoides R. Br., Protea odorata Thunb. and P. mucronifolia Salisb. All are
evergreen perennials. Protea odorata and P. mucronifolia grow as 0,5—0,8 m
high, single-stemmed, firm woody shrubs (caespitose nanophanerophytes). The
Serruria species grow as shrublets (chamaephytes) arising from a small lignotuber
at ground level as a number of sparsely branched lax stems to a height of
0,2—0,7 m, lowest for the sprawling S. trilopha.
* Present address: School of Environmental and Life Sciences, Murdoch University,
Murdoch, Western Australia.
Accepted for publication 16th March, 1978.
363
364 Journal of South African Botany
Methods were similar to those used by Milewski (1978). The thoroughness of
the investigation was severely limited by the study species’ relict status. All known
remaining populations were investigated, ““‘populations’’ of a species being arbi-
trarily defined as stands farther than 2 km apart (Milewski & Esterhuysen, 1977;
Milewski, 1978). Data were recorded from two populations of all species except
S. furcellata and S. cyanoides (only one population of each evidently remains),
and S. linearis (three populations). The substrate, and the physiognomy and
general floristics of vegetation, associated with each population were subjectively
recorded. All associate species in the Proteaceae and Restionaceae (Milewski,
1978), growing within 0,5 m of the study species, were listed. This was done for
each of a minimum total of 50 study plants scattered throughout each population
except for S. furcellata, of which only 25 plants could be found.
BiGaale
The western Cape coastal flats, showing distribution of Protea odorata (O), P. muc-
ronifolia (M), Serruria trilopha (T), S. linearis (L), S. furcellata (F) and S. cyanoides (C)
in relation to biotic areas. Localities were plotted using a twelfth-degree grid. Each symbol
represents the occurrence of the species within a square corresponding to one-ninth of the
area of a quarter-degree grid-square (smaller versions of the symbols were used where two
species were recorded in the same grid-square). See Milewski and Esterhuysen (1977) and
Milewski (1977) for explanation of biotic areas.
Habitat of threatened species of Serruria and Protea 365
RESULTS AND DISCUSSION
Serruria trilopha, S. linearis and S. furcellata are associated with adjacent
areas of pale, leached Quaternary sand (Table 1, Figs 1 & 2). This sand occurs as
a mantle of variable depth on flat to gently sloping landscapes (Table 1). The
associated vegetation is Coastal Fynbos (Table 1).
Herbarium labels indicate that S. cyanoides formerly occurred at altitudes
between 30 m and 550 m “‘on hill slopes and on flats’’. Collection sites include
“‘dunes’’ at the lowest altitudes and “‘sandy south-east slopes’’ of mountains at the
highest altitudes. Serruria cyanoides is today known only from aeolian sand
covering a gently sloping saddle on the mid-altitude slopes of a coastal mountain
(Table 1). The associated Coastal Fynbos differs from that associated with S.
apa) mage
sand — > clay
Fic. 2.
he habitats of 16 threatened species of Proteaceae and
Restionaceae on the western coastal flats of the south-western Cape excluding the Beaver
Valley, drawn subjectively on the basis of data on associates in the Proteaceae an
Diagrammatic representation of t
Restionaceae, and observation of substrates. 1 = Serruria trilopha , 2= S. Biase =
S. linearis, 4 = S. furcellata, 5 = Chondropetalum acockit, 6 = ciliata, 7 = egg
prominens, 8 = Restio micans, 9 = Leucadendron levisanus , IO = L. EAD z
Restio sabulosus, 12 = E. verreauxii, 13 = Protea mucronifolia, 14 = Diastella buekii,
= Protea odorata and 16 = L. verticillatum. Overlapping areas indicate that He ee
coexist in part of their habitat as associates (Milewski & Esterhuysen Pe aos “le
1978). Thamnochortus fraternus, a species of vee neutral to alkaline ;
omitted.
366 Journal of South African Botany
trilopha, S. linearis and S. furcellata (Tables 1 & 2). The presence of Tham-
nochortus erectus and Leucadendron coniferum (Table 2) suggests that the
associated sand is younger and less leached than that on which the other Serruria
species grow (Taylor, 1969; Boucher, 1972; Milewski, 1977). This substrate is
usually found elsewhere as eroded sublittoral dunes (Taylor, 1969).
The communities of Restionaceae and Proteaceae associated with S. trilopha
and S. linearis were 46,1 % similar to each other, whereas they were respectively
only 20,3 % and 12,5 % similar to that associated with S. cyanoides. These figures
were calculated on the basis of the Czekanowski coefficient following Milewski
(1978). This supports the suggestion that the habitat of S. cyanoides is distinct
from those of the other study species of Serruria.
Protea odorata and P. mucronifolia are also allopatric (Fig. 1). Both occur on
clay-rich soil on flats and gentle slopes (Table 1, Fig. 2). The associated
vegetation is a form of Coastal Rhenosterbosveld (Table 1, Fig. 1). The associated
communities of Restionaceae and Proteaceae (Table 3) were 30,3% similar
between the two species, on the basis of the Czekanowski coefficient (following
Milewski, 1978). This suggests that the habitat of P. odorata is distinct from that
of P. mucronifolia (Milewski & Esterhuysen, 1977; Milewski,.1977) despite the
apparent similarity of the associated climate and substrate.
The habitats of S. trilopha, S. linearis and S. furcellata may be compared with
those of partly sympatric threatened species also occurring on sand (Milewski &
Esterhuysen, 1977; Milewski, 1977, 1978). These species, including the congener
S. ciliata, grow under conditions of poorer drainage than the present study species
(Fig. 2). The habitats of P. odorata and P. mucronifolia, compared in the same
way, are similar to that of Leucadendron verticillatum (Fig. 2).
ACKNOWLEDGEMENTS
I wish to thank Professor E. A. C. L. E. Schelpe for the use of facilities at the
Bolus Herbarium, University of Cape Town. The study was supervised by Dr A.
V. Hall, project leader of the Rare and Endangered Plant Species Research Group,
and supported by the National Programme for Environmental Sciences. Miss E.
Esterhuysen and Dr J. P. Rourke kindly assisted with identification of Re-
stionaceae and Proteaceae respectively. An initial herbarium survey of endemic
species was undertaken by Mrs M. Bates, Miss G. M. van Niekerk and Miss G.
M. Micklem. I am grateful to Mrs M. L. Jarman, Mrs D. Jones and Miss C.
Davidge for typing the manuscript.
REFERENCES
BOUCHER, C., 1972. The vegetation of the Cape Hangklip area. M.Sc. thesis. Cape Town:
University of Cape Town.
Mi EWSKI, A. V. and ESTERHUYSEN, E., 1977. Habitat of Restionaceae endemic to western
Cape coastal flats. JIS. Afr. Bot. 43: 233-241.
Habitat of threatened species of Serruria and Protea 367
MILEwskI, A. V., 1977. Habitat of Restionaceae endemic to the south-western Cape coastal
flats. JJ S. Afr. Bot. 43: 243-261.
MiLewskI, A. V., 1978. Habitat of threatened Proteaceae endemic to western Cape coastal
flats. JIS. Afr. Bot. 44: 55-65.
Tay or, H. C., 1969. Vegetation of the Cape of Good Hope Nature Reserve. M.Sc. thesis.
Cape Town: University of Cape Town.
Tay or, H. C., 1978. Phytogeography and ecology of Capensis. In: M. J. A. Werger (ed.),
Biogeography and Ecology of Southern Africa. The Hague: Junk.
Journal of South African Botany
368
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370 Journal of South African Botany
TABLE 2
Frequency of presence and absence of Restionaceous and Proteaceous associates, in populations of
study species of Serruria: 1 = S. trilopha, 2 = S. linearis, 3 = S. furcellata, 4 = S. cyanoides.
Numerator equals the number of populations in which the associate species was recorded within 0,5 m
of plants of the study species; denominator equals total number of investigated populations per study
species; ‘‘+’’ denotes associate species present in the same stand of vegetation as the study species but
not within 0,5 m. See Table 3 for explanation of percentages in parentheses.
Study species
Associate species
1 2 3
(4 %) (17 %) (O %)
Serruria decipiens R. Br. ..........-- Besvmenne lporeetteanthe 1/2 oP
Leucadendron cinereum (Solander ex Ait.) R. Br. ..... 1/2 ate
RrorealpulchrauRy croft eee nn aan 1/2 +
Leucadendron thymifolium (Salisb. ex Knight) Williams ar a5
Leucadendron lanigerum Buek ex Meisn. ...........- 1/2 1/3
Chondropetalum nudum (Nees) Rottb. ............... 1/2 1/3
SeruniaiburmanniiiRelB Laer arr ne 1/2 2/3
IRESTOUNONGTINUSEN (as eee rE 2/2 3/3
Leucospermum parile (Salisb. ex Knight) Sweet....... 2/3
RestiosformiofasiebermKcunthi een nee 2/3
INGO JUVOTBDS OR, 050000000009 00000000000000000 2/3
Cannomois acuminata (Thunb.) Pillans .............. 3/3
Leucadendron corymbosum Berg. ..............++-- ar
Restio)bifurcus) Neesiexs Mast.) 55 000 0h el leeee 1/3 ate
Staberoha cernua (L.f.) Dur. & Schinz .............. ay a
Chondropetalum acockii Pillans .................... aF ate
Hypodiscus aristatus (Thunb.) Nees................. ar
ESTO CR GUIGLATIES INES 2 000000000c000cc0000c0n06 oF
SATO GHITGE IR, VBR, ob ooo oncbaccccoosnoodneesees ar
RESID PA COSTE PMETS x o000005000000000000000000 oF
Elegiaijunceanly ew eer ee eee ene
Thamnochortus erectus (Thunb.) Mast. ..............
Ele pianyar inulatagNias eee
Leucadendron coniferum (L.) Meisn.................
Leucospermum conocarpodendron (L.) Buek ssp. vir-
idumBRourkeigee eee ee eee
WNDAT HARB WMD, 500000000¢00ancccgvvscc00
Leptocarpus vimineus (Rottb.) Pillans ............... 2/2 3/3 als
Thamnochortus punctatus Pillans ................... 2/2 3/3 [1
Staberoha distachya (Rottb.) Kunth................. 2/2 3/3 1/1
Leptocarpus impolitus (Kunth) Pillans ............... 2/2 2/3 ats
Thamnochortus obtusus Pillans..................... 2/2 2/3 1/1
WilldenowiavarescensiKuntheee eee eee 2/2 3/3 1/1
Restio sp. nov. aff. R. tenuissimus Kunth ............ 2/2 ar
Chondropetalum tectorum (L.) Pillans............... 1/2 ar
Rroteatrepensy (Ieo)ilemer vt ea ye en et nae 1/2
Willdenowiarstjiatalhunb ss nen nie 2/2 2/3 1/1
Leucospermum hypophyllocarpodendron (L.) Druce spp.
canaliculatum (Buek ex Meisn.) Rourke ........... 2/2 2/3 1/1
Diastella proteoides (L.) Druce .................... oF ay
WilldenowiaksulcataNiaste ene + 3/3 1/1
ET OLECHACA LO SEH Un bn 1/2 2/3
Leucadendron salignum Berg. ..................... 2/2 1/3 1/1
Protea scolymocephala Rech Bin Osteo Pek Sic, RS nat 2/2 2/3
MEO GISNUFTHIS MINN, scoccsoocconcconsecca.... 1/2 2/3 Ts
Hypodiscus willdenowia (Nees) Mast. ............... 2/2 2/3 1/1
Habitat of threatened species of Serruria and Protea 371
TABLE 3
Frequency of presence and absence of Restionaceous and Proteaceous associates in populations of
study species of Protea: 1 = P. odorata, 2 = P. mucronifolia. Explanation of occurrence values as for
Table 2. Percentages in parentheses at top of columns indicate proportion of associates unrecorded for
the other study congener.
; Study species
Associate species
1 2
(59 %) | (27 %)
lhamnocnoriusspachmanrniilsMasereeecee eee Cee ee eee OLE +
Pilidenowiarstriatamnunbs sarvnerae ee ick aeacia oes oie +
SCIUITONNGLAaSS ata BucksexeMeciSN ieee eerie ar
SiaDerONarGennuan (eats) POUT GA SChHINZ ise eerie re ceerociiee recietieiertice +
Willdenowiavarescensskunthice erences eee eters etscr tele orn +
Heptocarpus virmineus) (Rottb;)) Pillans). 32)) 2.0.) elles ele eee +
Hypodiscus willdenowia (Nees) Mast... .. 0.0... 5-22-2222 seer eee +
PIBRT: SYLATTONE INES, cocaacagboSo0csp 055 00000000000000400e0KGD000C 2/2
IannoChOrlusi{rullCOSUSIBELS we eee ee eee et 1/2
Thamnochortus punctatus Pillans .............0. 00sec settee teens 1/2
Leucadendron thymifolium (Salisb. ex Knight) Williams................+--- 1/2
Leucadendron verticillatum (Thunb.) Meisn. ..........--.+.+++eeese eee 1/2
Elegia sp. nov. Esterhuysen 34411 ............- 002 e eee eee eee eee eee 1/2
Restio sp. nov. Esterhuysen 30849 ..........--- see eee eee eee een ees 1/2
ihaninocnorntusopimususebillausieee eee e eee eC eer ere 1/2
Sara TPG TAT IMIS Teo no con cones ov0dsbucoss00G0GGDDODUDCCOSDOOUDN 1/2
Elegia vaginulata Mast. ..........--2 00 eee eee cece teet nent n tees 1/2
Restio sp. nov. Esterhuysen 32525 ..........-. 22s eee eee neers 1/2
Leucadendron cinereum (Solander ex Ait.) R. Br. ......------ sees eee 1/2
Protea scolymocephala Reich. ........-+-++ssscsser eee t rete teenies 1/2
Restio duthieae Pillans .-.......--+----- eee see eee tee ee 2/2 +
RESID TT? VEIN sccncoo soccc000 cp OD DDDACODDDND OH BGUOGOOBDONDDNNN 1/2 +
Hypodiscus paludosus Pillans .......-..+++++ 220s e seer etter trees 2/2 1/2
Chondropetalum rectum (Mast.) Pillans ......-.--+-++ss esses eset trees D2, 1/2
Leucadendron lanigerum Buek ex Meisn. ......----++++++200 reer ttt 1/2 1/2
Leptocarpus rigoratus Mast. .........+-+-:-+++ eset enter: 2/2 2/2
Restio cuspidatus Thunb. .-+.-.------- 0+ 22920077 2/2 2/2
Blegia parviflora Kunth ........-.--2 22-2022 2 208 2/2 2/2
Leucadendron corymbosum Berg. ...---+-+++:++seer retest ss 1/2 ae
IRESTOUTIPOTUSER OLD Seer elie eo hei + 8
Restio sp. nov. Esterhuysen 30859 ........---+---+- 2200s t cert A
Leucadendron stellare (Sims) Sweet .....-.--+++:sse teeter tre trrs sess AB
Thamnochortus sporadicus Pillans .....--+--+++++scesr ret trtr tests (+) i
SETFUTIG CANAICANS ROBE. oie ease eee iciel teeter Rec ce
JIS. Afr. Bot. 44 (4): 373-379 (1978)
FURTHER NOTES ON PROTEA IN SOUTH AFRICA
J. P. ROURKE
(Compton Herbarium, Kirstenbosch)
ABSTRACT
Three new species, Protea nubigena Rourke, P. piscina Rourke and P. inopina
Rourke, as well as two new subspecies, P. longiflora Lam. ssp. potbergensis Rourke
and P. roupelliae Meisn. ssp. hamiltonii Beard ex Rourke, are described.
UITTREKSEL
AANVULLENDE NOTAS OOR PROTEA IN SUID-AFRIKA
Drie nuwe spesies, Protea nubigena Rourke, P. piscina Rourke en P. inopina Rourke
asook twee subspesies, P. longiflora Lam. ssp. potbergensis Rourke en P. roupelliae
Meisn. ssp. hamiltonii Beard ex Rourke, word beskryf.
In concluding the groundwork towards completing a revision of Protea in
South Africa, it has been found necessary to describe certain additional taxa. This
is the final contribution in a series of miscellaneous notes published preparatory to
the appearance of a revision of the South African species of Protea.
Protea nubigena Rourke, sp. nov.; a habitu brevi ramosissimo, inflorescentiis
parvulis, stylis 35-40 mm longis, foliis oblanceolatis vel ellipticis glabris glauces-
centibus 40-60 mm longis 7-14 mm latis, distinguitur. Crescit scopuli graminei
verticales ad 2 300 m supra mare in Dracomontes.
Fruticulus ramosissimus, 0,5—0,7 m altus, caulibus a caudice exorientes.
Caules ramosissimi, 2—5 mm in diam., nudi, sed foliis aggregatis apices versus;
rami cicatricibus foliorum delapsorum notati. Folia ascendentia, oblanceolata vel
elliptica, 40-60 mm longa, 7-13 mm lata, glauca, glabra; apices obtusi. [nflores-
centia globosa, 40—50 mm in diam. Receptaculum conicum, acutum, 10 mm
latum, 10-12 mm altum. Bracteae involucrales 3—4 seriatae, eburneae vel roseae
soridae, glabrae; series intima oblonga—linearia, 25-30 mm longa, 3—7 mm lata,
apices rotundatos concavos; series externa lata ovata 10-15 mm longa, 10 mm
lata, margines brunescentes. Perianthium 30-35 mm longum, tubus glaber extus
sed marginibus interioribus distalibus setosis; ungues et limbi setosi sparsi,
indumento fulvo. Stylus 35-40 mm longus, arcuatus adaxialis, albus. Stigma
peranguste linearie, 10 mm longum roseum.
An erect, much branched shrublet 0,5—0,7 m in height, to 0,5 m in diam., with
numerous 10-15 mm diam. stems arising from a subterranean caudex, very
Accepted for publication ist June, 1978.
373
374 Journal of South African Botany
occasionally with a short single main stem to 20 mm in diam., swollen at base to
form a bulbous caudex. Stems highly branched, woody, 2-5 mm in diam., with
short (20-50 mm) growth increments; glabrous, covered with dark verrucose bark;
leaf scars prominent; bare of leaves except for ultimate and penultimate growth
increments. Leaves erect, ascending, oblanceolate—elliptic, glaucous, glabrous,
40-60 mm long, 7-13 mm wide; apices obtuse, margins distinct. Inflorescence
globose, 40-50 mm in diam., surrounded by several axillary shoots. Receptacle
conic, acute, 10 mm wide 10-12 mm in height. Involucral bracts 3—4 seriate,
greenish cream to dirty pink, glabrous; inner series linear-oblong 25-30 mm long,
3-7 mm wide, apices rounded, slightly concave; outer series broadly ovate 10—15
mm long, 10 mm wide, margins brown. Perianth 30-35 mm long, slightly
adaxially curved; tube region 10 mm long, glabrous externally but inner margins
densely setose distally; claws sparsely setose with a ferruginous indumentum on
inner and outer surface; limbs 10 mm long, apices sharply acuminate, outer
surface sparsely setose with a tawny ferruginous indumentum. Anthers 4, subses-
sile, 5-6 mm long, filaments 1,5 mm long. Style 35-40 mm long, adaxially
curved, tapering subterminally, white. Pollen presenter very narrowly linear,
simple, scarcely differentiated from style, 10 mm long, pink. Ovary obconic 5—6
mm long, covered with long, straight, tawny trichomes. Hypogynous scales
lanceolate, acuminate, 1-5 mm long.
Type: Royal Natal National Park above Policeman’s Helmet Ridge on vertical cliffs of
south-east aspect, facing Tugela Gorge, on the ridge leading to The Witches, 2 300 m (7 600
ft), 4/4/1978, J. P. Rourke 1635 (NBG, holo.!; PRE, NU, J, BOL, K, MO, S and in herb.
Royal National Park Mont-aux-Sources, iso.).
Protea nubigena may be distinguished from all other Protea species of the
summer rainfall region of South Africa, on account of its dwarf, highly branched
growth habit, the very small inflorescences and flowers (styles 35-40 mm long),
and the small, oblanceolate, glabrous, glaucescent leaves 40-60 mm long, 7-14
mm wide.
A most remarkable species presently known from a single locality in the Natal
Drakensberg within the Royal Natal National Park where it grows on vertical cliffs
and ledges at an elevation of 2 300 m (7 600 ft), on Basalt lavas of the Stormberg
series. The associated vegetation consists of coarse turf composed of Heteropogon
contortus and Themeda triandra with Buchenroedera lotononoides, Cliffortia
browniana, Helichrysum sutherlandii and Talbotia elegans being the other pre-
dominant species on the ledges.
SPECIMENS EXAMINED:
NATAL—2828 (Bethlehem): Natal National Park, Policeman’s Helmet Ridge,
on cliff, south aspect (-DB), 8 000 ft., 18/7/1964, Esterhuysen 30249 (BOL);
Royal Natal National Park, Policeman’s Helmet Ridge, Oct. 1976, J. Scotcher s.n.
Further notes on Protea in South Africa 375)
(NBG /10, 236 ); Royal Natal National Park, above Policeman’s Helmet Ridge on
vertical cliffs of south-east aspect facing Tugela Gorge, on the ridge leading to The
Witches, 7 600 ft., 4/4/1978, Rourke 1635 (NBG, PRE, NU, J, BOL, K, MO, S).
Protea piscina Rourke, sp. nov.; fruticulus nanus caespitosus; foliis scabris,
acicularibus—canaliculatis vel linearibus, 15-30 cm longis, 2-5 mm latis, bracteis
luteis vel subroseis indumento albo sericeo, distinguitur.
Fruticulus nanus caespitosus, 1 m in diam., 300 mm altus. Caules ramosis-
simi, subterranei, 3-8 mm in diam., glabri, folia in caespites terminales gerentes.
Folia acicularia—canaliculata vel linearia—plana, 150-300 mm longa, 2-5 mm
lata, scabra, pubescentia dum immatura sed mox glabra; apices cuspidati. Inflores-
centiae obconicae—turbinatae, 35-50 mm longae, 40-50 mm in diam. Recep-
taculum late conicum, depressum, 5—15 mm latum, 5—10 mm altum. Bracteae
involucrales 4—5 seriatae, luteae vel subroseae, indumentum sericeum, album;
series externa ovata acuta, 5-7 mm lata, 7-10 mm longa; series intima,
oblonga—linearia, 20-35 mm longa, 2—5 mm lata, apices rotundatos concavos.
Perianthium 30-35 mm longum; tubus glaber, quadrangulatus, 8-10 mm longus;
Ungues dense lanati; limbi lineares acuminati, lanati. Stylus 30-35 mm longus,
arcuatus adaxialis. Stigma peranguste lineare, 5-7 mm longum.
Dwarf, tufted shrublet, forming clumps or mats to | m in diam., 300 mm in
height. Stems subterranean, highly divaricate, 3-8 mm in diam., glabrous,
producing terminal tufts of leaves at ground level; new shoots regenerate from
subterranean stems after burning. Leaves acicular—canaliculate to linear and flat,
150-300 mm long, 2-5 mm wide; scabrous, margins horny, apices sharply
cuspidate to uncinate; pubescent when young, soon glabrous but occasionally with
some pubescence remaining, particularly in the petiolar region. /nflorescences
obconic, turbinate, 35—50 mm long, 40-50 mm in diam. Involucral receptacle
broadly conic, depressed, 5-15 mm wide, 5-10 mm high. Involucral bracts 4—5
seriate, pale yellow, occasionally flushed with pink; outer series ovate acute, 5—7
mm wide, 7-10 mm long, margins ciliate; inner series oblong—tinear, 20-35
mm long, 2-5 mm wide, apices concave, rounded; covered with a soft, white,
sparsely sericeous indumentum. Perianth 30-35 mm long, straight to slightly
adaxially arcuate; tube region glabrous, quadrangular, winged, 8-10 mm long,
inner surface slightly pubescent distally; claws slender, densely lanate with a
shaggy white indumentum, occasionally interspersed with tawny trichomes; limbs
10 mm long, linear, markedly acuminate, with a white lanate indumentum. Style
slightly adaxially arcuate, tapering terminally, 30-35 mm long, occasionally
puberulous proximally. Pollen presenter simple, linear, acute, scarcely differen-
tiated from style, S—7 mm long. Ovary 3-5 mm long, obconic, covered with long,
straight tawny trichomes. Hypogynous scales lanceolate acute, 2 mm long.
Type: Upper Olifants River Valley, between Visgat and Bollemakiesiekloof, Dec.,
Rourke 1506 (NBG, holotype; BOL, PRE, K, MO, S, M, isotypes).
376 Journal of South African Botany
Although clearly allied to P. scabra R. Br., which is characterised by the
presence of a rusty brown indumentum on the involucral bracts, P. piscina is
distinguished by the pale yellow (very rarely pink-flushed) involucral bracts
covered with a soft white, sericeous, indumentum, and the acicular—canaliculate
to linear leaves, 150-300 mm long, 2-5 mm wide. At no point do the areas of
distribution of the two species overlap. P. piscina is distributed to the north and
east of the range of P. scabra; from the Cedarberg to the Cold Bokkeveld and
Piketberg and with the extreme easterly populations occurring from Swellendam
along the southern foothills of the Langeberg, to Plattekloof.
SPECIMENS EX AMINED
CAPE PROVINCE—3218 (Clanwilliam): Piketberg, top of Versveld’s Pass
(-DC), March, Paterson-Jones s.n. (NBG 93932); Piketberg mountain, southeast
of town, Dec., H. C. Taylor 4479 (NBG, STE); Piketberg mountain (-DD), Feb.,
van der Westhuizen s.n. (BOL); Kafferskloof Farm Jan., Summers s.n. (NBG
85502).
3219 (Wuppertal): Grootberg, Clanwilliam, southwest slopes (-AC), Dec.,
Esterhuysen 4191 (BOL); top of Olifants river mountains, behind Warmbaths
(-CA), Sept., Stephens 6910 (BOL); Skoongesig, (-CC), Nov., Hanekom 530
(PRE); Grootfontein, Olifantsriver valley, Sept., Oliver 3986 (STE); hills a mile
southeast of Keerom, Nov., Pillans 8775 (BOL).
—3319 (Worcester); Plateau of Skurfdeberg between Olifants and Kromme
river (-AA), Dec., Andreae 197 (PRE, STE); road between Rosendalfontein and
Visgat, Nov., Pillans 9642 (BOL); Olifants river valley, Onderboskloof, Jan.,
Esterhuysen 14296 (BOL); Upper Olifants river valley between Visgat and
Bollemakiesiekloof, Dec., Rourke 1506 (NBG, BOL, PRE, K, MO, S, M); top of
Gydo Pass (-AB), July, Markétter s.n. (STE 19382); Witzenberg, top and eastern
slopes (-AC), Dec., Andreae 155 (PRE).
——3420 (Bredasdorp): Swellendam (-AB), June, Harold Fry s.n. 4985 in herb.
Galpin (BOL, SAM, PRE); hill close to Swellendam just north of town, Oct.,
Marloth 7957 (PRE); Langeberg, near Swellendam, Sept., Muir 2752 & 2753 (PRE);
hill southeast of Swellendam on road to Bontebok Park, Nov., Williams 1727
(NBG); below the Crown mountain, Swellendam, Aug., Wurts 270 (NBG); hills at
Voormansbosch (-BA), Sept., Zeyher 3671 (SAM); In lat. mont. pr. Swellendam,
Niven s.n. (K); Bontebok Park, Feb., Grobler 601 (STE).
eo at (Montagu): Buffelsjagtsrivier at Sparrbosch (-DC), 30/7/1831, Drége
(P).
——3321 (Ladismith): Plattekloof, near Riversdale (-CC), Oct., Dekenah s.n.
(NBG 95694).
Protea inopina Rourke, sp. noy.; a P. arborea Houtt., bracteis dense fer-
rugineis velutinis et habitu humilis, caulibus simplicibus (raro ramosis) ad 1 m
altis caudice exorienti, distinguendus.
Further notes on Protea in South Africa 3
Frutex humilis, ad 1 m altus. Caules simplices (raro ramosi), 10-12 mm in
diam., glabri, caudice exorienti. Folia elliptica vel late oblanceolata 120-180 mm
longa, 25-50 mm lata, glabra, glauca, cartilaginea vel coriacea. Bractea 5-7
seriatae, dense velutinae ferruginiae, praeter seriem intimam partialem pubescen-
tiem. Perianthium 80—90 mm longum; tubus glaber, 10 mm longus, porcatus;
ungues tenues, glabri; limbi 30 mm longi, crassi, tomentosi vel lanati. Stylus
80—90 mm longus, leviter arcuatus. Stigma 15—20 mm longum, tenue. Ovarium
obovoideum, 3—5 mm longum, trichomatibus fulvis vestitis.
Low, multiple-stemmed shrub 0,5 to 1 m in height, the stems arising from a
woody subterranean rootstock. Stems erect, glabrous, 10—12 mm in diam., simple
to occasionally branched, more usually simple. Leaves elliptic to broadly oblan-
ceolate, 120-180 mm long, 25—50 mm wide, glabrous, glaucous, cartilaginous to
coriaceous in texture, sessile, but frequently twisted in petiolar region. Inflores-
cence globose to spherical in bud, becoming elongate before anthesis with the
perianths projecting 30-50 mm beyond the involucral bracts prior to anthesis,
becoming ovoid when fully open, 100-120 mm in diam.; substipitate to stipitate,
stipe to 30 mm long. Jnvolucral receptacle prominent, conic-acute, 40-45 mm
wide, 20-30 mm high. Involucral bracts 5-7 seriate, covered with a densely
velutinous, ferruginous indumentum in bud stage; outer series ovate acute, tightly
imbricate, 10-15 mm wide, 10—20 mm long, covered with a densely velutinous
ferruginous indumentum; inner series broadly ovate to ovate—oblong, obtuse,
15-25 mm wide, 20-25 mm long, with a ferruginous indumentum towards apex
and midrib, otherwise glabrescent, pale green. Perianth 80-90 mm long, tube 10
mm long, prominently ridged, glabrous but minutely puberulous along inner
margin of tube lip; claws slender, glabrous; limbs stout, 30 mm long with a
densely tomentose to lanate white indumentum, except on the adaxial limb, apices
incurved very acuminate, densely lanate. Anthers 4, linear, subsessile, 20 mm
long, apical boss green, sagittate. Style 80-90 mm long, adaxially arcuate,
glabrous, tapering terminally. Pollen presenter 15-20 mm long, slender, linear,
not clearly differentiated from style. Ovary obovoid, 3-5 mm long covered with a
small tuft of straight tawny trichomes. Hypogynous scales 1,5 mm long, narrowly
and obliquely ovate. Fruits large, ovoid 10-12 mm long 6-7 mm wide, nut-like.
i ins i i-se-Berg and Pre-
Type: Bergvlei mountains, east of Paleisheuwel, between Bergvlei-se g
toriuskrans, above the farm Mieliedraai, 27/9/1977, J. P. Rourke 1619 (NBG, holo.!, PRE,
STE, BOL, MO, K, S, L, iso.!).
P. inopina is distinguished by the densely ferruginous, velutinous pubescence
on the involucral bracts (particularly in the bud stages) and by the dwarf growth
habit with simple (only very rarely branched) stems up to 1 m in height arising
ther significant diagnostic feature of this species is
f istent rootstock. Ano
ah eae 10-12 mm long and 6-8
the inordinately large size of the fruits. Ovoid in form,
378 Journal of South African Batany
mm in diameter they are three times the size of the fruits in P. arborea. Together
with those of P. magnifica, they are the largest fruits recorded in Protea.
SPECIMENS EXAMINED
CAPE—3218 (Clanwilliam): Bergvlei Mountains, east of Paleisheuwel, be-
tween Bergvlei-Se-Berg and Pretoriuskrans, above the farm Mieliedraai (-DB),
27/9/1977, Rourke 1619 (NBG, PRE, STE, K, BOL, MO, S, L); on the farm
Wilgerbosdrif, Olifants Rivier Mountains, about 13 miles northwest of Pieken-
nierskloof Pass, M. Schonken s.n. (STE 30939); Citrusdal area, 10/11/1966, M.
Vogts s.n. (NBG 85255 ).
Protea longiflora Lam. ssp. potbergensis Rourke, ssp. nov.
Frutex vel arbor parva, ad 4 m alta. Folia ovata vel late ovata, basibus
cordatis; velutina, postea glabrescentia; 60-90 mm longa, 45—65 mm lata, sed
decrescentia (35 mm longa, 25 mm lata), ad apicem caulis. Stylus 110-130 mm
longus.
Shrub or small tree to 4 m. Leaves ovate to broadly ovate, cordate at base,
velutinous at first becoming glabrous about a year later, margins thickly ciliate;
lower leaves 60-90 mm long, 45-65 mm wide, decreasing in size towards shoot
apex to 35 mm long and 25 mm wide. Style 110-130 mm long.
Type: Potberg, south slopes near Hamerkop, flowering in hort. J. P. Rourke 25/5/1976,
Rourke 273 (NBG, holo.; PRE, BOL, K, MO, S, iso.)
Apparently confined to the Potberg and adjacent coastal flats, this subspecies is
distinguished by its large, ovate to broadly ovate leaves, velutinous initially with
thickly ciliate margins.
SPECIMENS EXAMINED
CAPE—3420 (Bredasdorp): Potberg, south slopes near Hamerkop (-BC), fid. in
hort. J. P. Rourke 25/5/1976, Rourke 273 (NBG, PRE, BOL, K, MO, S); Mr
Michlers farm, south end of Potberg, Dec. (in fruit), Williams 629 (NBG); Middle
south slope of the Potberg, margin of a stream, Oct. 1940, Pillans 9382 (BOL).
Protea roupelliae Meisn. ssp. hamiltonii Beard ex Rourke, ssp. nov.; a habito
nano prostrato, distinguitur.
Fruticulus procumbens, maxime divaricatus ad 300 mm altus, teges caespitosa
ad 1 m in diam. formans; caule principali singulari. Rami crassi, horizontaliter
effusi. Inflorescentia et flos characteres ut in sub-speciem typicam.
_Low, procumbent, highly divaricate shrublet to 300 mm in height, forming a
tufted mat to 1 m in diam.; with a single main stem. Branches stout, short,
sprawling horizontally along ground. Inflorescence and floral characters as in
typical subspecies.
Further notes on Protea in South Africa 379
Type: Wartberg, Nelshoogte Forest Reserve, 30/1/1975, Rourke 1467 (NBG, holo.;
PRE, K, MO, iso:).
This extraordinary local race of P. roupelliae was discovered by Dr P. D.
Hamilton of Barberton early in 1957. He showed the populations to Dr J. S. Beard
who proposed that it be named P. roupelliae var. hamiltonii, although no formal
description was ever published. It appears that Beard referred to this local race as
P. roupelliae var. hamiltonii in correspondence with the Botanical Research
Institute, Pretoria, but he did not affix a manuscript name to any herbarium
specimens. Moreover, for several years plants have been offered for sale by a
well-known firm of Protea growers in Cape Town under this nomen nudum.
Beard’s proposed epithet is here adopted and validated although it is considered
that subspecific rank is more appropriate for this dwarf race of P. roupelliae on
account of its morphological and geographical distinctness.
SPECIMENS EX AMINED
TRANSVAAL—2530 (Lydenburg): Nelshoogte Plateau (-DD), Dec., P. D.
Hamilton s.n. No. 1525 in herb. Beard (NU); Nelshoogte Plateau, Aug., P. D.
Hamilton s.n. No. 1024 in herb. Beard (NU); Nelshoogte, a small creeping plant,
P. D. Hamilton s.n. No. 985a, in herb. Beard (NU); Nelshoogte Plateau, March,
J. S. Beard 880 (NU); Dr. Hamilton Nature Reserve, Nelshoogte (-DB), Dec.,
Muller 2127 (PRE); Dr Hamilton Protea Reserve, Jambila Forest Reserve, Nov.,
Edwards 4127 (PRE); Wartberg, Nelshoogte Forest Reserve, Jan., Rourke 1467
(NBG, PRE, K, MO).
ACKNOWLEDGEMENTS
I am very much indebted to the Director of the Natal Parks, Game and Fish
Preservation Board for generously placing the full facilities of his organisation at
my disposal to facilitate the collection of P. nubigena. In particular, I am deeply
grateful to a member of his staff, Mr John Scotcher, whose sustained interest over
several years led to the rediscovery of this species’ natural habitat and without
whose planning and organisation the mission would not have been the success it
was. Finally, were it not for the sturdy assistance of Mr R. E. Levitt and Mr J.
Scotcher, who supported the author on a rope while the type material was being
collected, an adequate account of P. nubigena would not have been possible.
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JIS. Afr. Bot. 44 (4): 381-386 (1978)
FLAVONOIDS OF THE SUMMER RAINFALL PROTEAS AND THEIR
CHEMOTAXONOMIC SIGNIFICANCE
C. W. GLENNIE
(Department of Chemistry, University of the Witwatersrand, Johannesburg)
L. E. DAVIDSON
(Department of Botany, University of the Witwatersrand, Johannesburg)
ABSTRACT
Flavonoids were surveyed (by paper chromatography) in leaves of 15 summer rainfall
Protea species. Five different 3—O—glycosides of quercetin were found, viz. the arabinoside,
galactoside, glucoside, rhamnoside and rutinoside. Most species contained only 3 different
glycosides but one species, P. laetans L. E. Davidson, contained all five. P. laetans, also,
was the only species to contain leucoanthocyanins. All species, except three contained
abutin and none contained cyanogenic compounds.
UITTREKSEL
FLAVONOIEDE VAN DIE SOMERREENSTREEK PROTEAS EN HULLE CHEMIES-
TAKSONOMIESE BETEKENIS
’n Papierchromatografiese opname is gedoen van die voorkoms van flavonoiede in die
blare van 15 Protea spesies van die somerreénstreek. Vyf verskillende 3—0—glikosiede van
kwersetien is gevind, nl. die arabinosied, galaktosied, glukosied, rhamnosied, en rutinosied.
Meeste spesies het slegs drie van die glikosiede bevat. P. laetans daarenteen al vyf. P.
laetans was ook die enigste spesies wat leukoantosianiene bevat het. Al die spesies behalwe
drie het arbutien bevat, maar geen sianogeniese stowwe nie.
INTRODUCTION
Comparatively little is known about the phytochemistry of indigenous South
African plants. A study of the summer rainfall proteas was therefore carried out to
determine their flavonoid glycoside content. This would be of value for several
reasons. Mainly, it would serve to determine the chemical relationships between
the various species in this group and also variations within species. A survey of
flavonoids and other phenolic compounds has not yet been carried out and it
would be worthwhile to screen for novel compounds.
Another reason for a chemical survey of these plants is the position of Protea
laetans. This species was recently described by Davidson (1975) and it was
desirable to examine its flavonoid content and compare this with related taxa.
Also, P. caffra, P. multibracteata, and P. rhodantha all have very similar
morphology and it would be of interest to know whether this similarity carried
through to their flavonoid content.
—————
Accepted for publication 19th May, 1978.
381
382 Journal of South African Botany
Flavonoids were chosen as the compounds of interest in this study because of
their many desirable properties as summarised by Harborne (1967). Elsworth and
Martin (1971) reported the possible presence of quercetin in four species reported
here and also the possibility that kaempferol could be present in trace amounts.
Since their work was done on acid-hydrolysed plant extracts it was decided to
expand on this and attempt to isolate and identify all the unhydrolysed flavonoid
glycosides for this study.
In another study, members of the Proteaceae were found to contain both
hydroquinone and its glucosylated derivative arbutin. These compounds were
reported to be present in all of the 29 species studied (van Oudtshoorn, 1963). To
extend this survey, all plants in the present study were examined for the presence
of arbutin. Recent work in this laboratory has established the presence of acylated
forms of arbutin in several Protea spp. (Perold and Rosenberg).
METHODS
The plant material was collected from a variety of locations in South Africa
and air dried. Voucher specimens are deposited in the Moss Herbarium, University
of the Witwatersrand. The dried leaves were milled and extracted with 80%
methanol. After concentration, the extract was applied to Whatman 3 MM paper;
the flavonoids were separated and purified using n-butanol:acetic acid: water
(4:1:5) (BAW), 15% acetic acid, and n-butanol:ethanol:water (4:1:2,2) (BEW)
(Harborne, 1967). The flavonoid glycosides were compared with authentic sam-
ples by R; values (Table 1) in BAW, phenol:water (500:125) (PhOH), 15%
HOAc and water. The u v spectra of all five glycosides were similar, Band I
absorbing at 258 nm and Band II absorbing at 362 nm in 80% methanol.
For comparison purposes two-dimensional paper chromatograms were run on
Whatman No. 1 paper, using BAW and 15% HOAc. When viewed under uv light
it was difficult to locate all the dark absorbing flavonoids amongst the blue
fluorescent compounds. This was overcome by placing a fluorescent sheet of paper
under the chromatogram which blended with the blue spots thus clearly showing
the dark flavonoids.
All samples of plant material were examined for hydrogen cyanide-producing
compounds in the following manner. A small sample of each aqueous plant extract
was hydrolysed with B-glucosidase which was obtained commercially from sweet
almond meat. The released HCN was detected by picric acid papers suspended
over the solution in an enclosed round bottom flask. All positive tests were
checked by the Dilute Hydrochloric Acid Test (Vogel, 1942).
The presence of leucoanthocyanins was determined by hydrolysing the plant
extracts with 2N HCl in a test tube and then extracting with amyl alcohol.
Delphinidin and cyanidin were identified by comparison with authentic standards
in respect to R; values on paper chromatograms and u v maxima (Harbone, 1973).
Flavonoids of Summer Rainfall Proteas 383
Unhydrolysed, methanolic plant extracts were used for the survey of arbutin
and hydroquinone. Spots of these extracts were applied to Whatman No. 1 paper
chromatograms which were developed overnight in BAW or BPW
(n-butanol:pyridine:water-6:4:3). After air drying, the chromatograms were
sprayed with Pauly’s Reagent (0,5 % solution of diazotised sulfanilic acid in 10%
aqueous Na,CO;). The arbutin and hydroquinone appeared as red spots on a
white or pale cream background.
TABLE 1.
R,; Values of Phenolic Glycosides of Summer Rainfall Protea Species.
R,; values (x100) in
Compound 15%
PhOH HOAc H,O
Quercetin 3-arabinoside ............ 61 33 07
Quercetin 3-galactoside ............ 62 48 12
Quercetin 3-glucoside.............. 62 35 06
Quercetin 3-rhamnoside ............ 56 5) 22
Quercetin’ 3-rutinoside .-........... 46 58 27
/MMONIN" £ 6. cob 0 BOO OR ae aan Coes 65
87
RESULTS AND DISCUSSION
The flavonoids in the leaves of twenty-six different samples of fifteen species
of Protea are reported in Table 2. Along with the chemical results are also shown
the geographical location from which the plant sample was collected. All plants
contained only the one aglycone, quercetin, and any variations arose from the
different 3-0-glycosides present. Flavonols are the flavonoids generally associated
with the more primitive woody members of the Dicotyledoneae. The presence of
the flavonol quercetin in this group of proteas suggests that it is rather primitive.
Also, since it is the only aglycone present and has a simple glycosylation pattern,
all members of this group are probably of a similar evolutionary age.
Quercetin has been reported previously in four of the species studied here
(Elsworth & Martin, 1971; van Oudtshoorn, 1963). The presence of kaempferol
was reported in two of these species but in this study no kaempferol could be
found. The plants showed four different patterns of quercetin glycoside distribu-
384 Journal of South African Botany
tion. The galactoside, glucoside and rhamnoside were distributed throughout while
the arabinoside and rutinoside were restricted in their distribution.
P. multibracteata and P. rhodantha showed chemical variation within the
species while all other species had only one pattern of flavonoids. Four samples of
multibracteata contained three of the quercetin glycosides while one sample
contained an extra glycoside. P. rhodantha exhibited three different glycoside
patterns. As can be seen from Table 2 P. multibracteata and P. rhodantha are the
only two species examined which show the property of chemical variation.
It is probable that the rutin (quercetin-3-0-rhamnoglucoside) found in these
plants is biosynthesized from quercetin-3-0-glucoside by the addition of rhamnose.
This sequence of building up diglycosides of flavonoids from the monoglycosides
has been suggested by Harborne (1975).
All plants in this study were tested for cyanogenesis and found negative.
Hegnauer (1969) reported that P. cynaroides L., one of the winter rainfall Protea
species, gave a positive test for HCN. This is the only member of the sub-family
Proteoideae which was reported positive. (All the South African Proteaceae except
for one genus, Brabeium, belong to this sub-family.) The present study yielded
contrary results, for when material of P. cynaroides was collected from three
different sources, it was found negative in all cases. On the other hand, ten of
thirty-seven species of Leucadendron and Leucospermum tested for cyanogensis
gave positive results (Glennie).
The only species which contained leucoanthocyanins was P. laetans. The
anthocyanins released on acid hydrolysis of the plant material were identified as
cyanidin and delphinidin. In addition to being the only species to contain
leucoanthocyanins, P. laetans is the only species to contain all five quercetin
glycosides. These two properties appear to be unique to P. laetans and delimit this
new species from the others.
All plant samples of summer rainfall proteas, with the exception of P.
rubropilosa, P. nubigena and P. comptonii, contained arbutin and many contained
its aglycone hydroquinone. It is probable that the hydroquinone came from the
breakdown of arbutin during the collecting and drying of the plant material. For
this reason the distribution pattern of hydroquinone is not considered, as it could
be of dubious value. Including this study, arbutin has now been reported present in
41 species of 6 genera in the Proteoideae. Outside this sub-family and the
Ericaceae, where it is widespread, its occurrence is rare and erratic, being found
mainly in Malus and Pyrus spp.
Arbutin was found to be absent from P. comptonii and P. rubropilosa and to
be replaced by several esters of 2-hydroxy-4-hydroxymethylphenylalloside. Both
6-0-cinnamate and 6-0-benzoate of 2-hydroxy-4-hydroxymethylphenyl B-D-
allopyranoside were found in these two species (Perold, Beylis and Howard,
1973b). The newly described species, P. nubigena, also does not contain arbutin
but neither does it contain the same replacement compounds as P. rubropilosa
Flavonoids of Summer Rainfall Proteas 385
TABLE 2
Distribution of Phenol Glycosides in Summer Rainfall Protea Species, by paper chromato-
graphic analysis.
Species Compound Geographical Source
LEO BAGS Sees 6)
P. caffra Meissner [Far oe + Magaliesburg
P. comptonii Beard qeege dk Haenertsburg
P. curvata N.E.Br. ap Spe gp) aa at Barberton
P. dracomontana Beard + + 4+ op Drakensberg
P. gaguedi J. F. Gmelin + + 4+ + Barberton (2 samples)
P. laetans L. E. Davidson a sr Gp Se gk ok Blyde River (2 samples)
P. multibracteata Phillips | i oa + Oribi, S. Coast Natal
P. multibracteata Phillips oF OE + Port St Johns (2 samples)
P. multibracteata Phillips + ff + Kokstad
P. multibracteata Phillips + + + + + S. Coast Natal
P. nubigena Rourke i ot Tugela Gorge
P. parvula Beard ae ap Dullstroom
P. rhodantha Hook f. + + + + Nels Hoogte
P. rhodantha Hook f. oF SP AP + Blyde River
P. rhodantha Hook f. + + + + Long Tom Pass
P. rhodantha Hook f. apap ab + Iron Crown
P. rhodantha Hook f. cs oar oar OE + Iron Crown
P. rhodantha Hook f. + + + + + Blyde River
P. rhodantha Hook f. 7 ooP ap ap Sb Haenertsburg
P. rhodantha Hook f. + + + + + Sheba (2 samples)
P. rubropilosa Beard a oP Oop Blyde River
P. roupelliae (dwarf form)
Meissner qr ap + Barberton
P. simplex Phillips + + S. Coast Natal
P. transvalensis Phillips Se tect + Haenertsburg
P. transvalensis Phillips te oar ae + Iron Crown
P. welwitschii Engel FOP ar RP Johannesburg
Key 1 — Quercetin-3-0-arabinoside, 2 — Quercetin-3-0-galactoside,
3 — Quercertin-3-0-glucoside, 4 — Quercetin-3-0-rhamnoside,
5 — Quercetin-3-0-rutinoside, 6 — Arbutin.
and P. comptonii. In P. nubigena arbutin appears to be replaced by several
phenolic compounds which are yet to be identified.
It is of interest that P. comptonii and P. rubropilosa lack arbutin and that both
contain the replacement glycosides, rubropilosin and pilorubrosin. They also
contain the same complement of flavonoids, viz. the galactoside, glucoside and
thamnoside of quercetin. These species have similar morphology which tends to
distinguish them from related taxa and one of the features separating them from
each other is their geophysical distribution.
P. comptonii is found in a small area south of Barberton, Transvaal, while IP,
rubropilosa is found on the escarpment of the north-eastern Transvaal. Previously,
the 6-0-benzoate of 2 hydroxy-4-hydroxymethylphenyl—D-gluco-pyranoside had
been isolated from the winter rainfall Protea lacticolor Salisb. (Perold, Beylis and
386 Journal of South African Botany
Howard, 1973a). It appears that this ester also replaces arbutin in P. lacticolor as
no arbutin or hydroquinone could be detected.
Two different samples of P. gaguedi were collected and examined. Both of the
plants examined have similar morphology and inhabit the same ecological zone.
The main difference between these samples is their time of blooming; one blooms
in August while the other blooms in October. This time of flowering appears to be
the only difference as both plants exhibited identical flavonoid and arbutin content
as well as cyanogenic properties.
ACKNOWLEDGEMENT
Professor G. W. Perold (Department of Chemistry) is thanked for his encour-
agement and for his support of this work. Dr. J. P. Rourke (National Botanic
Gardens of South Africa, Kirstenbosch) is thanked for supplying a sample of
Protea nubigena.
REFERENCES
Davipson, L. E., 1975. A New species of Protea in the Transvaal. JI S. Afr. Bot. 41:
207-210.
EtswortH, J. F. and Martin, K. R., 1971. Flavonoids of the Proteaceae, Part 1. A
Chemical Contribution to Studies on the Evolutionary Relationships in the S.
African Proteoideae. JI S. Afr. Bot. 37: 199-212.
GLENNIE, C. W. Unpublished results.
Harpsorne, J. B., 1967. Comparative Biochemistry of the Flavonoids. London and New
York: Academic Press.
HarBorne, J. B., 1973. Phytochemical Methods. London: Chapman and Hall.
HARBORNE, J. B., 1975. Flavone and Flavonol Glycosides. In: Harborne, J. B., Mabry, T. J.
and Mabry, Helga (eds.), The Flavonoids. London: Chapman and Hall.
HEGNAUER, R., 1969. Chemotaxonomie der Pflanzen. Volume 5. Basel and Stuttgart:
Birkhauser.
PEROLD, G. W., BEYLis, P. and Howarp, A. S., 1973a. J. chem. Soc. Perkin 1: 1973,
638-643.
PEROLD, G. W., BEYLis, P. and Howarp, A. S., 1973b. J. chem. Soc. Perkin 1: 1973,
643-649.
PEROLD, G. W. and ROSENBERG, M. E. K. Unpublished results.
VAN RHEEDE VAN OupTSHOORN, M. C. B., 1963. Distribution of phenolic compounds in
some South African Proteaceae. Planta med. 11: 399-406.
VoGEL, A. I., 1942. Qualitative Chemical Analysis. London: Longmans, Green.
IS. Afr. Bot. 44 (4): 387-390 (1978)
CHROMOSOME NUMBERS IN LACHENALIA (LILIACEAE)
ROBERT ORNDUFF AND PaTRICIA J. WATTERS
(Department of Botany, University of California, Berkeley, California, U.S.A.)
ABSTRACT
__ Chromosome counts are reported for forty-one collections of Lachenalia, representing
sixteen species and numerous additional undetermined collections, ones of uncertain status,
or undescribed species. Chromosome numbers of n = Sy, I, ©, 1M; 1, 11H, 14, 1G, 118
22 and 28 were obtained. Chromosomal races are recorded in some species. For
Lachenalia, it is probable that x = 5, 6, 7, 8 and 9.
UITTREKSEL
CHROMOSOOMGETALLE IN LACHENALIA (LILIACEAE)
Chromosoomgetalle in 41 versamelings van Lachenalia word vermeld. Dit sluit 16
soorte en baie ander ongeidentifiseerde versamelings, soorte van onsekere status en
onbeskryfde soorte in. Getalle van n = 5, 6, 7, 8, 9, 10, 11, 12, 14, 16, 18, 22, en 28 is
verkry. Chromosomaal-rasse is aangetref by sekere soorte. In Lachenalia is dit waarskynlik
dat x = 5, 6, 7, 8, en 9 is.
INTRODUCTION
Lachenalia is a genus of ca. 90 species (Barker, pers. comm.) endemic to
South Africa and South West Africa (Namibia); most species occur in the
south-western Cape Province in an area with a Mediterranean climate. Though the
lines between many species are clear cut, in others there are intergradations and
other morphological variation patterns that have led to taxonomic confusion
(Moffett, 1936). Two workers have made chromosomal studies of the genus
(Moffett, 1936; de Wet, 1957); their counts plus scattered counts made by other
workers have revealed chromosome numbers of 2n = 14, 16, 22, 26, 28, 32, 42,
and 56. During 1970-71, and in 1974, the first author collected Lachenalia in the
Cape Province, sent the bulbs to Berkeley, and initiated a chromosomal survey of
the genus. The results of this survey are reported herein.
MATERIAL AND METHODS
Floral buds of cultivated specimens were fixed in 3:1 glacial acetic acid:
absolute ethanol, transferred to 70 % ethanol after fixation, refrigerated, and
anthers removed, squashed in acetocarmine. The resultant preparations were
examined with a microscope and the number of chromosomes determined.
Accepted for publication 29th May, 1978.
387
388
Journal of South African Botany
TABLE 1.
Chromosome numbers in Lachenalia.
Species
aff. algoensis Schoenl.
aff. aloides (L.f.) Hort
bulbifera Cyrillo
contaminata Ait.
elegans Barker
aff. fistulosa Bak.
hirta Thunb.
juncifolia Bak.
longibracteata Phillips
mutabilis Sweet
orchioides (L.) Ait.
pallida Lindl.
pustulata Jacq.
purpureo-caerulea Jacq.
trichophylla Bak.
unicolor Jacq.
aff. unicolor Jacq.
unifolia Jacq.
violacea Jacq.
Undescribed species:
aff. hirta
Unidentified species:
“reciprocal translocation present
fragment present
COTODAMN
\© 00 0°
Author’s collection Previous reports
number and locality
7386:
7704:
7159:
7339:
7377:
7444:
7419:
TEMP
7449:
7229:
7342:
WNT
7511:
7095:
7431:
7072:
Wa ie
7344:
Caledon
Elgin
Strandfontein
Breérivier
betw. Worcester
and Villiersdorp
Bulshoek Dam
Pakhuis Pass
Pakhuis Pass
Pakhuis Pass
Caledon
Goudini
Ezelfontein
Op-Die-Berg
Mamre Rd. Station
Bulshoek Dam
Klawer
Clanwilliam
Ceres
2n
2n
2n
2n
24 (de Wet, 1957)
14, 56 (de Wet, 1957)
16 (de Wet, 1957);
16, 17 (Moffett, 1936)
DSi
7307:
7353:
Wye
JD22:
7429:
7130:
7421:
WAP
7163:
7368:
7515:
Kalbaskraal
Mamre
Paarl
Steenbergs Cove
Darling
Bidouw Valley
Kapteinskloof
Bidouw Valley
Aurora
Darling
Brandvlei Dam
Citrusdal 2n = 21, 22, 24, 26
(de Wet, 1957)
7439: Bulshoek Dam
7193:
7356:
Spektakel Pass
Worcester
8024:
8033:
7954:
7952:
7973:
Clanwilliam
Clanwilliam
Spektakel Pass
Okiep
betw. Vredendal
and Vanrhynsdorp
Pakhuis Pass
Bidouw Valley
N11 and Douse the
Glim Rd.
8044:
8051:
8036:
Chromosome numbers in Lachenalia (Liliaceae) 389
RESULTS
Forty-one collections were examined representing 16 species and several
undetermined collections, those of uncertain taxonomic status, and two unde-
scribed species. Chromosome numbers of 2n = 10, 12, 14, 16, 18, 22, 28, and 56
were obtained (Table 1).
DISCUSSION
It is clear that Lachenalia is chromosomally more diverse than previously
reported, since our counts of 2n = 10, 12, and 18 are new for the genus. Clearly
several base numbers exist: x = 5, 6, 7, 8, and 9. The high diploid numbers can be
considered polyploid ones based on various combinations of these base numbers,
though it is possible that higher, non-polyploid base numbers may also exist. Few
cytological aberrations were noted; a reciprocal translocation was noted in one
collection of L. mutabilis, and a fragment was observed in a collection of L.
pallida. For several taxa intraspecific polyploidy exists. Diploids and tetraploids
occur in L. orchioides; tetraploids and octoploids occur in L. elegans. In the latter,
populations with the two chromosome numbers occur very near each other. For
some species, reports by other workers indicate the existence of chromosomal
races at other levels. For example, two collections of L. mutabilis had n = 5,
whereas de Wet (1957) reported 2n = 14 and 56 for plants attributed to this
species. Likewise, in L. orchioides and its close relatives, n = 7 and 14 are
reported herein, whereas both de Wet (1957) and Moffett (1936) reported 2n =
16; the latter worker also recorded 2n = 17 for this species. For L. unifolia, n =
11 is reported from three widely separated populations; de Wet (1957) reported 2n
= 21, 22, 24, and 26 for this species.
Barker (unpub.) has recognized two provisional, major and as yet unnamed
groups within Lachenalia based on seed characters. All named taxa in Table 1
with n = 7, 14, or 28 fall into group 1 except for L. trichophylla, which has been
assigned to group 2. However, Barker (pers. comm.) states that the seed of L.
trichophylla is so distinctive that it and several other species with similar seeds
may be relegated to still a third group. However, we have obtained n = 5 for L.
mutabilis, also a member of group 1, although de Wet (1957) obtained numbers
for this species based on x = 7. All species we have examined with n = 8 and 11
fall into group 3.
It is thus clear that not only is Lachenalia a variable genus in its morphology,
but it is also unusually variable in chromosome numbers. The diverse chromosome
numbers reported for various species groups in this genus suggest particularly
problematical assemblages that merit further cytotaxonomic study.
ACKNOWLEDGMENTS
We are indebted to Dr M. S. Cave and Dr Peter Goldblatt for their assistance
with this project. These studies were supported in part by grants from the National
390 Journal of South African Botany
Science Foundation, the Bache Fund of the National Academy of Sciences (both
U.S.A.) and the C.S.I.R. We are particularly indebted to Miss W. F. Barker for
her continued interest in this project and for providing access to her considerable
store of knowledge regarding Lachenalia.
REFERENCES
DE WET, J. M. J., 1957. Chromosome numbers in the Scilleae. Cytologia 22: 145-159.
MoreettT, A. H., 1936. The cytology of Lachenalia. Cytologia 7: 490—498.
Ritey, H. P., 1962. Chromosome studies in some South African monocotyledons. Can. J.
Genet. Cytol. 4: 50—SS.
JIS. Afr. Bot. 44 (4): 391-418 (1978)
TEN NEW SPECIES OF LACHENALIA (LILIACEAE)
W. F. BARKER
(Bolus Herbarium, University of Cape Town)
ABSTRACT
Ten new species of Lachenalia are described, and the characters of the ripe seeds are
used for the first time as additional diagnostic features in the genus.
UITTREKSEL
TIEN NUWE LACHENALIA SOORTE (LILIACEAE)
Lien nuwe Lachenalia spesies word beskryf. Die eienskappe van die ryp sade word as
aanvullende diagnostiese kenmerke in die genus Lachenalia voorgestel en aangewend.
INTRODUCTION
Since the publication of Baker’s monograph on the genus Lachenalia in Flora
Capensis V1, 1896-1897, collections of many new taxa have lain dormant in
herbaria, particularly those of Rudolph Schlechter, and many others have been
collected in recent years, a number of them apparently, as far as is known, with a
very restricted distribution. It is proposed to publish them in sets of ten species, in
order to validate the names, to be used in the monograph which is in preparation.
Lachenalia peersii Marl. ex Barker, sp. nov.
Differt a L. rosea floribus brevioribus albis urceolatis basi rotundatis, segmen-
tis externis gibbosis viridofuscis, segmentis internis apicibus recurvis, staminibus
stylisque inclusis. Seminibus, globosis, arillo inflato, terminale, mensure medio ad
longo.
Plant up to 350 mm high, usually less. Bulb globose or depressed globose,
20-25 mm diam; outer tunics light brown, membranous. Leaves 1—2 lorate, up to
300 mm long and 10-25 mm broad, loosely clasping the base of the peduncle, or
sometimes with a clasping base up to 70 mm long, green or tinged with maroon
below, and sometimes above. Peduncle slender 50-200 mm long, usually dark
maroon. Inflorescence racemose, few to many flowered, up to 120 mm long and
20-25 mm diam. Flowers urceolate, rounded at the base, lax, spreading to
slightly cernuous, carnation-scented; pedicels 1,2 mm long; bracts narrow lanceo-
late, membranous; perianth 7-10 mm long; tube 2-3 m long, white; outer
segments ovate up to 6 mm long, white with greenish brown gibbosities, inner
segments obovate up to 8 mm long, with upper edges recurved, white, stamens
Accepted for publication 16th June, 1978.
391
392
Journal of South African Botany
Hermeanium AreicanumM BoLUBiANUM.
g A
rromarut! esd Sager
Lath unitio. farsii, thie oe
anus, Or. eke, ale tack of
fioyal Ket. L ftw L9LE. J lowed: *
Hit Loess gardens A ynteng. as.
Per hese anon / PLP Legit Viele Suars.
i 7
HOLE PEG Feexsié, pe
ee
Fic. |
Isotype sheet of Lachenalia peersii, in Bolus Herbarium No. 16360.
Ten new species of Lachenalia (Liliaceae) 393
included, as long as the outer segments, ovary globose 3 mm diam., green; style
up to 4 mm long. Capsule obovate, membranous, 6 mm diam; seeds globose; aril
inflated, terminal, narrow, medium to long.
DIAGNOSTIC CHARACTERS
Lachenalia peersii is allied to L. rosea Andr. but differs in having the
urceolate flowers rounded at the base, and the tips of the inner segments recurved.
It is also characterised by having a carnation scent.
|
imm
\
1 2
Fic. 2
Lachenalia peersii: 1. seed side view; 2. seed from above; 3. seed from below.
Type Material: CAPE PROVINCE—3419 (Caledon): Hermanus, (-AC), November
1915, Peers s.n. sub Herb. Marloth 7263 (PRE, holo.); November 1919 Peers s.n. sub.
BOL 1/6360 (BOL, iso.).
This attractive species was collected at Hermanus in 1912 by Mr V. S. Peers,
who grew it in his garden in Kenilworth, Cape. In November 1915 he gave
flowering specimens to Dr Rudolph Marloth, who preserved them in his herbarium
collection, and gave the species the manuscript name Lachenalia peersii in honour
of its collector. In December 1917 a photograph by Mr Peers, of a pot of bulbs in
flower, was published in ‘‘South African Gardening and Home Life’’, with the
caption ““Lachenalia peersii a new variety found in Hermanus and recently
named’’. In November 1919 further specimens from Mr Peers’ garden were
preserved in the Bolus Herbarium. The sheet was annotated in the hand of Mrs L
Bolus ‘‘Lachenalia peersii Marl. Typus in Litt’’ and it was presumed to be a valid
species. However as no trace of a description can be found in literature, Dr
Marloth’s manuscript name has now been validated.
He named it after the man whose name is usually associated with Peers Cave
at Fish Hoek, which he and his son Bertie excavated and made famous. It is not
generally known that Mr Peers was an enthusiastic and well informed botanical
collector of some note, who contributed as many as 700 collections of succulent
and bulbous plants to Kirstenbosch, in the early years after it was established.
394 Journal of South African Botany
The species is widely distributed in the south western part of the Caledon
district, where it occurs in rocky ground. Out in the open it is often rather stunted,
but plants flowering on recently burnt ground can become strong and robust.
SPECIMENS EXAMINED
CAPE PROVINCE—3418 (Simonstown): Palmiet River Bridge (-BD),
30/10/1962, Thomas s.n.sub. NBG 70017 (NBG); Sunny Seas, 29/9/1967, and
27/10/1967, Barker 10516 (NBG); Harold Porter Botanic Gardens, Bettys Bay,
27/10/1970, Ebersohn s.n.sub. NBG 92682 (NBG); Disa Kloof, Bettys Bay,
9/12/1966, (fruiting); 1/11/1968, (flowering) Rourke 686 (NBG); Hangklip,
8/12/1965, (fruiting) Barker 10368 (NBG); Waterfall Kloof, Hangklip Nov. 1948,
Stokoe s.n.sub. SAM 61313 (SAM).
3419 (Caledon): 12 m. W of Caledon (-AB), 17/9/1934, Salter 4861 (BM,
BOL, K); Hermanus (-AC), Nov. 1915, Peers s.n.sub. Herb. Marloth 7263
(PRE, holo); Nov. 1919, V. S. Peers s.n.sub. BOL 16360 (BOL); Nov. 1921,
Rogers s.n.sub. PRE 26566 (PRE); 8/11/1963, Thomas and Barnard s.n.sub.
NBG 70714 (NBG); Hermanuspetrusfontein, 23/10/1897, E. E. Galpin 4755
(PRE, K); Riviera Hotel Flats, 10/10/1916, Purcell 107 (SAM); 4/10/1918,
Purcell s.n.sub. (SAM); near Hermanus, 16/10/1966, Barker 10484 (NBG);
Voelklip Kloof, 18/11/1973, Esterhuysen 33357 (NBG); Voelklip Bot.Soc. Re-
serve, 22/10/1969, Rourke s.n.sub. NBG 93557 (NBG); Onrust River, 1/12/1951,
Esterhuysen 19268 (BOL); Shaws Mt, Sept. 1938, Gillett 4464 (BOL, K); De
Wets Bay, Kleinmond, 1/11/1968, Barker 10515 (NBG).
Lachenalia multifolia Barker, sp. nov.
Distinguitur foliis numerosis filiformibus teretibus pallide viridibus basi albis
succulentis, inflorescentia racemosa multiflora, pedicellis brevis patulis, floribus
campanulatis albis, segmentibus leviter cucullatis fere aequantibus pallide vir-
idibus gibbosis, staminibus perspicue exsertis rectis, seminibus globosis, arillo
jugoso terminale perbreve.
Plant 70-200 mm high, usually dwarf. Bulb globose 10-20 mm diam., outer
tunics papery, brownish. Leaves 5-13, usually as long or a little longer than the
peduncle, occasionally longer, 150-170 mm long; blade filiform terete, yellow-
green, broadening into white, fleshy, flattened bases 20-30 mm long and 5-8 mm
broad. Inflorescence racemose many-flowered up to 70 mm long and 30-50 mm
diam.; peduncle slender, green or tinged with red; pedicels spreading up to 10 mm
long, usually shorter. Flowers strongly scented, spreading; perianth 8 mm long;
tube 3 mm long, white; outer segments oblong 5,5 mm long, 3 mm wide, white
with pale green gibbosities; inner segments as long as outer, narrow oblong, white
with green keels, all segments slightly spreading with the apex slightly cucullate;
stamens exserted evenly spaced up to 7 mm long; ovary ovoid 3,5 mm long pale
Ten new species of Lachenalia (Liliaceae) 395
Fic. 3
1. flower side view; 2. flower from below;
outer perianth segment, 6. upper outer perianth
flower; 4. flower front view; 5. lateral
segment; 7. lateral inner perianth segment, 8. lower inner perianth segment; 9. stamen; 10.
gynoecium, all x 33,
Lachenalia multifolia: 3. transverse section of
396 Journal of South African Botany
green; style finally as long as stamens. Capsule obovate membranous 5 mm diam.,
1-2 seeds in each loculus; seed ovoid, aril very short, ridged terminal.
DIAGNOSTIC CHARACTERS
Lachenalia multifolia most closely resembles L. polyphylla but is distinguished
by having the swollen bases of the leaves white and smooth, the white flowers are
campanulate with the segments slightly spreading and cucullate, while those of
L. polyphylla are tinged maroon, urceolate, with the tips of the inner segments
recurved.
C | | ‘)
1 2 3
Fic. 4
Lachenalia multifolia: 1. seed side view; 2. seed from above; 3. seed from below.
Type Material: CAPE PROVINCE—3319 (Worcester): Karoo Poort (-BA), 26/9/1944,
Barker 3053 (NBG, holo.).
Lachenalia multifolia is one of only five species in the genus having many
leaves. It resembles L. polyphylla superficially but is very distinct when compared
in detail, and their habitats differ widely; it is found on rocky hillsides, in rock
pans and rock crevices, while L. polyphylla favours flat open ground. The distribu-
tion of the species too is widely separated, L. multifolia occurs in the Worcester,
Ceres, and Calvinia districts, while L. polyphylla is only known from the Malmes-
bury, Piketberg and Tulbagh districts. The two species differ considerably in the
characters of their seeds, L. multifolia has a seed with a very small ridged aril, that
of L. polyphylla is covered entirely by an irregular pattern of ridges.
SPECIMENS EXAMINED
CAPE PROVINCE—3119 (Calvinia): 10 m N. of Calvinia (-BD), 25/9/1962
Levyns (NBG).
3319 (Worcester): Ceres (-AD), 9/1925, Nielson s.n. sub. Hort. NBG
1008/25 (BOL); Ceres Karoo 10/1931 H. Herre s.n. sub Hort. Stell. Univ. Gdns
3844 (BOL); Karoo Poort (-BA) 26/9/1944, Barker 3053 (NBG holo); 22 m N. of
Karoo Poort, 19/9/1954, H. Hall 947 (BOL);
——3320 (Montagu): Constable, 4/10/1940 (-AD), Compton 9737 (NBG);
10/1940, Esterhuysen 5083 (BOL).
Ten new species of Lachenalia (Liliaceae) 397
2
Oy
ri
3
3
Té
2CM
dj Nw aed
Fic. 5
enae: 1. flower side view; 2. flower from below; 3. flower front view;
lateral outer perianth segment; 6. lateral inner perianth
t; 8. gynoecium; 9. transverse section of leaf, all x 3.
Lachenalia esterhuys
4. upper outer perianth segment; 5.
segment; 7. lower inner perianth segmen
398 Journal of South African Botany
Lachenalia esterhuysenae Barker, sp. nov.
Distinguitur foliis 1-2 longis linearibus teretibus venetis basi supra
canaliculatae, pedunculo longo gracile, inflorescentia laxo, pedicellis longis
patulis, floribus cremeis campanulatis cernuis, staminibus perspicue exsertis.
Seminibus ovoideis, arillo jugoso, terminale, minuto.
Plant 150-450 mm high. Bulb globose or depressed globose 12—18 mm diam.,
with soft brown outer tunics, not produced into a neck. Leaves 1-2, linear, terete,
with a shallow channel down the upper face toward the base, scarcely clasping the
base of the peduncle, erect with the support of surrounding vegetation, 140-360
mm long, up to 5 mm diam., blue-green, shading to reddish-brown near the base,
without markings. Peduncle up to 340 mm long, very slender, tinged or minutely
spotted with reddish-brown. Inflorescence racemose 40-110 mm long many-
flowered, sterile portion very short, flowers lax, cernuous; bracts very small, apex
narrow lanceolate, membranous, translucent; pedicels 4-10 mm long, very slen-
der, spreading, cream. Flowers 4—6 mm long, 3 mm diam. at base, cernuous,
campanulate, rounded at the base; tube 1 mm long, cream; outer segments 3—4
mm long, slightly spreading, cream with green gibbosities; inner segments slightly
longer than the outer up to 5 mm long, slightly spreading, cream with a greenish
blotch near the apex, upper edge of margin recurved with a small central cusp;
stamens well exserted up to 13 mm long; ovary ovate, up to 3 mm long, green;
style exserted up to 12 mm long. Capsule obovoid, membranous, 3 mm diam.
seeds shining, ovoid, aril ridged, terminal, minute.
DIAGNOSTIC CHARACTERS
Lachenalia esterhuysenae is easily recognized by its small, cernuous, cream
flowers with well exserted stamens, and long spreading pedicels, and by its very
long, blue-green, filiform, terete leaves.
C) e
1 2
Fic. 6
Lachenalia esterhuysenae: 1. seed side view; 2. seed from above; 3. seed from below.
Type Material: CAPE PROVINCE—3219 (Wuppertal): S. Cedarberg, Sneeuwberg
Area, above Bakleikraal (-AC), E. Esterhuysen 34149 (BOL, holo.; K, iso.; NBG, iso.;
MO, iso.; PRE, iso.; S, iso.).
This interesting high altitude species was first collected on 30th September
1940 in the Pakhuis Pass by Miss E. E. Esterhuysen, while on a journey to
Ten new species of Lachenalia (Liliaceae) 399
Calvinia to witness the total eclipse of the sun. Only one flowering bulb was
collected, which was later illustrated. A second small collection was made on 11th
December 1950, also by Miss Esterhuysen, at the northern end of the Cedarberg
Sneeuwkop, below the shale band at 1 200 m, extending its known range somewhat.
It was found again in the first locality on 19th October 1965, when Mrs M.
Thomas and Miss W. Barker were travelling through the Pakhuis Pass. The plants
were growing at about 900 m on a rocky slope, appearing through low fynbos, and
the peduncles and leaves were elongated to penetrate to the light. Bulbs were
grown and ripe seed was collected on 30th November 1965. On 11th December
1975 another extensive collection was made by Miss Esterhuysen, at the southern
end of the Cedarberg Sneeuwberg, above Bakleikraal, below the shale band at
1 200 m. It consisted of flowering and young fruiting plants some of which were
grown, and produced mature seeds in December 1976. It is named in honour of
Miss E. E. Esterhuysen, who was Botanical assistant at the Bolus Herbarium for
many years, and has always been most helpful and generous in supplying material
and information, particularly of high altitude species.
SPECIMENS EXAMINED
CAPE PROVINCE—3219 (Wuppertal): Pakhuis Pass, Clanwilliam, (-AA),
3/9/1940, E. Esterhuysen s.n.sub NBG 72233 (NBG); 19/10/1965, Barker 10319;
Cedarberg Sneeuwberg (-AC), 11/12/1950, E. Esterhuysen 18023 (BOL), S.
Cedarberg area above Bakleikraal, 11/12/1975, E. Esterhuysen 34149 (BOL,
holo.; K, iso.; NBG, iso.; MO, iso.; PRE, iso.; S, iso.).
Lachenalia congesta Barker, sp. nov.
Distinguitur foliis 2 ovatis oblongis prostratis supre atroviridis infra mar-
roninos, inflorescentia spicato congesto, pedunculo brevissimo saepe non suprater-
raneo, floribus cremeis gibbosis virido-fuscis, segmentibus internis pellucidis apice
recurvatis, staminibus inclusis, seminibus ovoideus, arillo jugoso terminale, per-
breve, testa Toveolata.
Plant 80-140 mm high, dwarf. Bulb obovate, 15-24 mm diam. with light
brown, spongy outer tunics produced into a long neck up to 40 mm long;
leaf-sheath obtuse, transparent, extending above the neck. Leaves 2, resembling
those of Massonia; blade thick, ovate to oblong, subacute, prostrate 35-75 mm
long and 34-56 mm wide, dark green or tinged with maroon, sometimes with a
few irregular blotches above, suffused with maroon below, margin maroon; bases
white, clasping the peduncle. Inflorescence a very dense spike 25 mm diam., up to
70 mm long, flowers suberect to spreading, strongly scented, peduncle very short
often none above ground. Bracts narrow-lanceolate, up to 5 mm long membran-
ous, white. Flowers sessile up to 13 mm long; tube 3 mm long, very pale blue;
outer segments 7 mm long, ovate erect, white shading to pale yellow with green
400 Journal of South African Botany
G.J. Lewis.
Lachenalia con ; oe
gesta: 1. flower front vi 5) i i
Bas, CEE Uo WOME view X 3; 2. flower side view X 2; 3. flow
4 aie Bac section of flower X 2; 5. outer perianth segment and tae Say
perianth segment and stamen X 3; 7. gynoecium X 3; Del G. J. Lewis
Ten new species of Lachenalia (Liliaceae) 401
gibbosities and pellucid recurved apices; inner segments 10 mm long, narrow
oblong emarginate, cream shading to yellow with green keels, pellucid apices
recurved; stamens included, 5 mm long, filaments white; ovary oblong up to 4 mm
long, green; style usually short 2-3 mm long, white. Capsule obovate, carpels
each with a portion of the persistent style forming a cusp at the apex; seed
comparatively large, globose with a much reduced conical ribbed aril, the surface
apparently with a depressed pattern. (Only one capsule and one seed was
produced).
DIAGNOSTIC CHARACTERS
Lachenalia congesta is a very distinctive species, with its dense inflorescence
and prostrate leaves. It belongs to the group of plants which have seeds with the
aril almost completely reduced. It appears to be most closely allied to L. isopetala,
another very distinct species, as their seeds are similar in shape and the largest in
the group.
Fic. 8
Lachenalia congesta: 1. seed side view; 2. seed from above; 3. seed from below.
Type Material: CAPE PROVINCE—3220 (Sutherland): Sutherland (-BC), 8/7/1968, F.
Stayner s.n. sub. NBG 93575 (NBG, holo.); 18/6/1970, Stayner s.n. sub. NBG 93574
(NBG, iso.).
The earliest record of this unusual and interesting species with its congested
inflorescence emerging from two prostrate, broad, dark green leaves, was made by
Mr P. Ross-Frames, who collected it at Muldersfontein 10 miles west of
Middelpos, in the Calvinia district, on 7th August 1933. He brought it to the
Bolus Herbarium, where Miss G. J. Lewis made the accompanying painting. She
noted that the leaves lying flat on the ground resembled those of a Massonia.
When not in flower they could very well be mistaken for one. About 20 years later
it was collected S.S.E. of Sutherland by Acocks, and in 1955 by Leistner. On the
8th of August 1968 another collection, this time in bud, was brought to the
Compton Herbarium by Mr F. Stayner, for many years Curator of the Worcester
Karoo Botanic Garden, also collected at Sutherland. Bulbs flowered in a pot on the
22nd of July 1968 and again on the 18th June 1970. Only one capsule matured,
402 Journal of South African Botany
and one seed developed, which had a very reduced aril, and the testa appeared to
have a depressed pattern on it. The plant according to Acocks grows in shale
gravel, in mountain Rhenosterveld.
SPECIMENS EXAMINED
CAPE PROVINCE—3120 (Williston): near Muldersfontein, 10 m W. of
Middelpos (-CC), 7/8/1933, P. Ross-Frames s.n. sub. BOL 2/365 (BOL).
3220 (Sutherland): Sutherland (-BC), 8/7/1968, Stayner s.n.sub. NBG 93575
(NBG, holo.); 18/6/1970, Stayner s.n. sub. NBG 93574 (NBG, iso.); 14 m S.S.E.
of Sutherland (-BD), 24/8/1953, Acocks 16935 (K); 19,5 m S.S.E. of Sutherland,
1/9/1955, Leistner 281 (K).
Lachenalia namaquensis Schltr. ex Barker, sp. nov.
Distinguitur foliis 1-2 linearibus lanceolatis canaliculatis ad apicem
semiteretibus, inflorescentia spicata, floribus rectis vel patulis, periantho externo
urceolato tubo caeruleo segmentis magenteis, segmentibus internis longioribus
patentibus recurvatis supernis albis inferiore magenteo, staminibus segmentibus
internis brevioribus, seminibus globosis, arillo jugoso, terminale, breve.
Plant up to 230 mm high. Bulb globose, small 10-15 mm diam.; outer tunics
thin, membranous, russet-brown, produced into a short neck; bulbils produced at
soil level from a few stolons from the base. Leaves 1-2, linear-lanceolate, up to
140 mm long; clasping base up to 60 mm long white, blade green, conduplicate or
with inrolled margins, narrowing to the semiterete apex. Peduncle slender up to
120 mm long. Inflorescence spicate up to 100 mm long, 20 mm diam., few to
many flowered; bracts very small, subobtuse. Flower suberect to spreading;
perianth 8-11 mm long, narrow urceolate; tube 2-3 mm long, bright blue; outer
segments 5 mm long, ovate, subacute, magenta with maroon-purple gibbosities;
inner segments up to 9 mm long, oblanceolate, narrowing to the base, the two
upper overlapping, spreading at the apex, shining white, tinged magenta at the
apex; lower segment narrower, deep magenta, apex recurving; stamens a little
shorter than the inner segments; ovary ovate 2-3 mm diam. style up to 6 mm
long. Capsule oblong, membranous, 5 mm diam.; seeds globose, aril ridged,
short, terminal.
DIAGNOSTIC CHARACTERS
Lachenalia namaquensis is characterised by its one to two leaves, which have
linear lanceolate, conduplicate blades, narrowing to a semiterete apex. The flowers
are sessile, narrow urceolate, usually blue at the base, shading to magenta, the
Inner segments are longer than the outer, the upper two are white while the lower
one is deep magenta. (A colour form with rose coloured flowers was found at
Klipfontein by Mr J. W. Mathews on 27th October 1930.) The bulb produces
Ten new species of Lachenalia (Liliaceae) 403
2CM
Fic. 9
side view; 2. flower from below; 3. transverse section of
flower; 4. flower front view; 5. lateral outer perianth segment; 6. upper outer perianth
Lachenalia namaquensis: 1. flower
segment; 7. lateral inner perianth segment, 8. lower inner perianth segment; 9. gynoecium,
all x 3
404 Journal of South African Botany
1 2 3
Fic. 10
Lachenalia namaquensis: 1. seed side view; 2. seed from above; 3. seed from below.
bulbils at ground level, on long stolons from the base of the bulb, and the plants
are usually found in colonies.
Type Material: CAPE PROVINCE—2917 (Springbok): 6 m S. of Steinkopf, (-BD),
23/8/1959, Barker 9020 (NBG, holo.).
Lachenalia namaquensis was collected by Rudolph Schlechter as early as 23rd
September 1897, and he distributed it to many herbaria under its manuscript
name. As its distribution appears to be confined to Namaqualand, the name is
appropriate and it is now validated.
It is most closely allied to L. framesii Barker which differs from it in having
shorter yellow flowers, with all the inner segments tipped with deep magenta, and
the leaves are lanceolate with undulate margins. The seeds in the two species are
similar.
In its natural habitat L. namaquensis is a dainty dwarf species, which increases
rapidly by means of bulbils on long stolons, and it usually occurs in colonies. In
1959 it was found growing in mass, in pockets on huge granite boulders near
Steinkopf, and more recently in stony clay soil, or in sandy soil, in other localities.
SPECIMENS EXAMINED
CAPE PROVINCE—2816/—2817 (Oranjemund/Vioolsdrif): Richtersveld,
26/8/1931, Herre s.n.sub. Hort. SUG 3862 (BOL); 9/9/1936 Herre s.n.sub. Hort.
SUG 3902 (BOL).
——2917 (Springbok): Steinkopf-Klipfontein (-BA), 26/8/1931, Herre s.n.sub.
Hort. SUG 3836 (BOL); Steinkopf-Obob, Sept. 1935, Herre s.n.sub. Hort SUG
3856 (BOL); 4 m N.W. by W. of Steinkopf, 21/9/1957, Acocks 19529 (NBG);
Klipfontein, Oct. 1930 and 1931, Mathews s.n.sub. Hort. NBG 2228/29 (BOL),
Oct. (colour var.); Oct. 1926, Pillans 5712 (BOL); Steinkopf (-BA/-BO),
23/9/1897, R. Schlechter 11383 (B, BOL, G, GRA, K, LD, PRE, 7L)\, SEVER).
Salter 3755 (BOL); Steinkopf-Platteklip, Herre s.n.sub Hort SUG 3862 (BOL); 1
m W. of Steinkopf, 19/10/1971 and 19/10/1973, Hall 4175 (NBG), 26/8/1931,
Herre s.n.sub Hort SUG 3837 (BOL); 6 m S. of Steinkopf (-BD), 23/8/1959,
Barker 9020 (NBG, holo.); Spektakel Mt. (-DA), 26/9/1974, Goldblatt 2788
Ten new species of Lachenalia (Liliaceae) 405
(NBG); 14 m W. by S. of Springbok, 24/9/1957, Acocks 19571 (NBG); 4 m E. of
Springbok (-DB), 15/9/1948, Salter s.n.sub. NBG Hort. 2075/1933 (NBG).
Lachenalia verticillata Barker, sp. nov.
Distinguitur folio unico glauco-viridi falcati vel lanceolati conduplicati infra
maculati basi amplectenti fasciata, inflorescentia spicata floribus patentibus in
quoque verticillo tres, periantho externo urceolato tubo caeruleo segmentibus
pallidis, segmentibus internis longioribus patulis recurvatis apice magenteis, stam-
ninbus segmentibus internis brevioribus, seminibus globosis arillo jugoso ter-
minale mensure medio.
Plant up to 250 mm high. Bulb globose up to 20 mm diam., outer tunics
spongy, brown; basal sheath membranous, obtuse, up to 20 mm long. Leaf 1,
falcate to lanceolate, conduplicate; blade spreading, up to 90 mm long and 20 mm
broad, blue-green, blotched with purple on the lower side, margin undulate;
clasping base up to 50 mm long, banded with purple above shading to magenta at
the base. Peduncle slender up to 90 mm long. Inflorescence spicate, up to 100
mm long and 20—25 mm diam.; flowers in verticils of three, spreading; bracts very
small. Flowers 9-12 mm long, urceolate; tube 3 mm long dull blue; outer
segments paler, ovate convex, subacute, 6 mm long and 5 mm broad; lateral inner
segments obovate oblong, 10 mm long, 3 mm broad near the apex, with purple
recurved tips; lower inner segment slightly longer and conduplicate with a purple
recurved tip; stamens included, 6 mm long; ovary globose, 2 mm long; style 6 mm
long. Capsule globose 6 mm diam.; seed globose aril ridged, terminal, medium
length.
DIAGNOSTIC CHARACTERS
Lachenalia verticillata is distinguished by its single falcate to lanceolate
conduplicate leaf which is spotted on the lower side of the blade, banded on the
clasping base and the sessile flowers are arranged in three-flowered verticils. The
outer perianth is urceolate and broadest at the base, and the inner segments are
longer, spreading, recurved, and tipped with purple.
Fic. 12
1. seed side view; 2. seed from above; 3. seed from below.
Lachenalia verticillata:
406 Journal of South African Botany
2CM
Fic. 11
Lachenalia verticillata: 1. flower side view; 2. flower from below; 3. flower front view; 4.
transverse section of flower; 5. lateral outer perianth segment; 6. upper outer perianth
segment; 7. gynoecium, X 3.
Ten new species of Lachenalia (Liliaceae) 407
Type Material: CAPE PROVINCE—2917 (Springbok): 3 m S.E. of Steinkopf, 18/9/193
Salter 3746 (BOL. holo.; BM, iso.; K. iso.). opf, 18/9/1933,
Only two collections have been made of this unique species, in spite of the fact
that it flowers in September when many people visit the area in a good flower
season. It was first collected by H. Bolus near O’Okiep in September 1833. The
second collection was made fifty years later, in September 1933 by Paymaster
Captain T. M. Salter who found it not very far from the original locality, at
Springbok, in Namaqualand.
SPECIMENS EXAMINED
CAPE PROVINCE—2917 (Springbok): 3 miles S.E. of Springbok (-BA/-BC),
18/9/1933, Salter 3746 (BOL, holo.; BM, iso.; K, iso.); Near O’Okiep (-DB),
9/1883, H. Bolus 6590 (BOL, K, SAM).
Lachenalia concordiana Schltr. ex Barker, sp. nov.
Distinguitur folio unico, lamina angusta lineari conduplicato curvato, basi
amplectenti viridi fasciata, floribus sessilibus late campanulatis, staminibus seg-
mentibus internis aequantibus, seminibus oblongis, arillo jugoso, terminale, men-
sure medio.
Plant up to 200 mm high. Bulb obovate, 12-20 mm diam.; with many dark
brown tunics produced into a neck up to 25 mm long. Leaf 1, up to 120 mm long;
blade narrow linear-lanceolate, conduplicate, spreading, curved, green up to 80 mm
long; clasping base slender, banded with green in the upper half, up to 70 mm
long. Peduncle short, up to 40 mm long above the clasping base. Inflorescence
spicate 20-100 mm long, up to 20 mm diam., few to many flowered, flowers
spreading, and usually arranged in three-flowered verticils. Flowers widely cam-
panulate 9-12 mm long; tube 3 mm long (colour not recorded); outer segments 7
mm long ovate acute spreading slightly, tips recurved; inner segments 9 mm long,
the two lateral obovate, subacute overlapping, recurving at the apex; lower ovate
acute spreading to form a lip; stamens as long as the inner segments; ovary ovoid
4 mm long; style finally longer than the stamens. Capsule membranous 8 mm
long, ovate; seed oblong, aril ridged, terminal, of medium length.
DIAGNOSTIC CHARACTERS
Lachenalia concordiana is distinguished by its single leaf with a narrow linear
lanceolate, conduplicate blade, and the clasping base banded with green. The
spicate inflorescence has the widely campanulate flowers arranged in three-flowered
verticils.
408 Journal of South African Botany
Plantae Schlechterianae Austro-Africanae. |
liter secundum.
{
CUCATEL SE 1A 1
Regio occidentalis
Fic. 13
Sheet of Lachenalia concordiana, herbarium of the British Museum, Natural History,
London.
Ten new species of Lachenalia (Liliaceae) 409
1 2 3
Fic. 14
Lachenalia concordiana: 1. seed side view; 2. seed from above; 3. seed from below.
Type Material: CAPE PROVINCE—2917 (Springbok): 15 m N.E. of Springbok (-DB),
8/9/1950, Barker 6762 (NBG, holo., iso.).
Until recently only two collections of this distinctive species were known. The
original one, collected on 19th September 1897 by R. Schlechter, at Concordia in
Namaqualand, was distributed to many herbaria under his manuscript name, now
validated. The second was made by W. F. Barker on 8th September 1950 near
Springbok. The discovery of a third collection among the incertae in the Bolus
Herbarium, collected by C. L. Leipoldt in September 1941 near Garies, a
considerable distance to the south, indicates that the species has a wider distribu-
tion than was at first thought. Fortunately this material includes capsules with ripe
seeds, which were absent in the earlier collections; these assist in placing the
species in the group with ridged, terminal arils.
SPECIMENS EX AMINED
CAPE PROVINCE—2917 (Springbok): Concordia (-DB), 19/9/1897,
Schlechter 11322 (B, BM, G, GRA, K, LD, Z); 19/9/1897, Schlechter 11321?
(PRE); 15 m N.E. of Springbok, 8/9/1950, Barker 6762 (NBG holo., iso.).
—3018 (Khamiesberg): near Garies (-CA), Sept., 1941, G. L. Leipoldt 3575
(BOL).
Lachenalia glaucophylla Barker, sp. nov.
Distinguitur folio unico glauco anguste lanceolato conduplicato patentis vel
curvatis, basi amplectenti, inflorescentis racemosa angusta multiflora, floribus
patulis cremeis late campanulatis, segmentibus fere aequantibus, staminibus rectis
perspicue exsertis. seminibus oblongis, arillo jugoso terminale stature medio vel
breve, in carina angusta decurrente prolongata.
Plant 90-250 mm high, usually dwarf. Bulb globose 10—20 mm diam., outer
tunics spongy, brown, disintegrating, produced into a short neck. Leaf usually 1,
410 Journal of South African Botany
very occasionally 2, narrow-lanceolate, conduplicate; blade up to 130 mm long,
usually less, up to 14 mm at ground level, spreading or curved almost at right
angles, glaucous green without markings; clasping base up to 70 mm long, white,
surrounded by a membranous transparent sheath up to half its length. Peduncle
usually very short, occasionally up to 60 mm long above ground level, tinged with
brown. Inflorescence racemose, narrow, 20 mm diam., up to 80 mm long,
many-flowered, flowers spreading; bracts minute, obtuse, membranous; pedicels
2-3 mm long, spreading, cream. Flowers very small, up to 5 mm long; tube 0,5
mm long, cream; outer segments 4 mm long, ovate, spreading, cream with green
gibbosities; inner segments 5 mm long, narrow oblong, spreading, cream with
green keel; stamens exserted 7 mm long, spreading, filaments cream; ovary
globose 1—2 mm diam., green; style 3—5 mm long. Capsule globose 4 mm diam.,
membranous, about 2 seeds to each loculus; seeds oblong, comparatively large,
shining, aril terminal, short or medium, extending as a narrow ridge down to the
ridged micropyle.
Fic. 15 a, b.
a. two plants, one in bud; b. inflorescence close up; c. plants in
bud.
Lachenalia glaucophylla:
Ten new species of Lachenalia (Liliaceae) 411
DIAGNOSTIC CHARACTERS
Lachenalia glaucophylla is easily distinguished by its 1 (or very occasionally
2) linear-lanceolate conduplicate, curved, glaucous leaf, its dense narrow racemose
inflorescence composed of very small cream flowers, with spreading segments of
almost equal length, and much exserted spreading stamens. Its seed is ovoid, with
a terminal, short or medium ribbed aril.
Lachenalia glaucophylla: 1. seed side view; 2. seed from above; 3. seed from below.
Type Material: CAPE PROVINCE—3319 (Calvinia): 11 miles from Downes on
Klipwerf road (-BD), 14/10/1974, M. Thomas s.n. sub. NBG 105737 (NBG, holo.).
The earliest collection of this dainty new species to be traced, was collected by
Dr. Rudolph Marloth in October, 1917 at Leliefontein in Namaqualand, and is
412 Journal of South African Botany
preserved in the National Herbarium, Pretoria. It consists of two very small
specimens in the young fruiting stage, which made its identification difficult. More
recent collections representing all stages, including good flowering material have
been made, and it is now possible to decide that they represent a previously
undescribed species, which is distinctive in leaf and floral characters. The seeds
belong to the large group with ribbed arils.
It was found growing in colonies on open flats with sparse vegetation, in very
shallow pans where silt had collected. The recent collections have all been made
in the Calvinia district, some distance from the Marloth specimens, thus extending
the known distribution of the species considerably. The fact that it flowers in
October may account for the fact that it has not been more widely collected.
SPECIMENS EXAMINED
CAPE PROVINCE—3018 (Kamiesberg): Kamiesbergen, Leliefontein (-AC),
9/1917, Marloth 8101 (young fruits) (PRE).
— 3319 (Calvinia): Moordenaarspoort, 26 miles N.E. of Calvinia near Agter
Hantamsberg (-BD), 6/10/1953, G. J. Lewis 2630 (SAM); 11 miles from Downes
on Klipwerf road, 14/10/1974, M. Thomas s.n. sub. NBG 105737 (NBG, holo.).
3120 (Williston): 40 m E. of Calvinia on Williston road (-AC), 13/10/1976,
Barker 10554 (young fruit) (NBG).
Lachenalia sargeantii Barker, sp. nov.
Distinguitur foliis 2 linearibus lanceolatis conduplicatis patentibus, inflorescen-
tia racemosa, pedunculo atro-marronino, pedicellis longis patulis magenteis,
floribus cremeis pendulis tubulosis basi rotundatis magenteis suffusus, segmentis
externis gibbosis viridofuscis, segmentis internis longioribus apice macula virido-
fusca, staminibus inclusis, seminibus globosis, arillo membranoso terminale breve
in carina decurrente lato prolongata, testa colliculata.
Plant 200-300 mm high. Bulb 15-20 mm diam. with dark brown leathery
tunics, producing a nest of bulbils round the base of the bulb. Leaves 2,
linear-lanceolate, conduplicate, loosely clasping the base of the peduncle, suberect
to spreading, up to 300 mm long and 15 mm broad, yellow-green to bluish-green,
without markings. Peduncle 140-280 mm high, much longer than the raceme, dark
maroon and faintly blotched and lightly glaucous. Inflorescence racemose, 6—25
flowered; flowers pendulous concentrated into a head, sterile portion at apex very
short; bracts white or tinged with magenta, ovate lanceolate, short up to 4 mm
long; pedicels to lower flowers shorter, about 4 mm long, increasing in length
upwards to 15 mm long spreading to recurved, magenta. Flowers up to 25 mm
long, pendulous, tubular, rounded at the base and slightly constricted about 3 mm
above the base; tube 3-4 mm long; outer segments up to 15 mm long, oblong
lanceolate, cream with greenish-brown gibbosities; inner segments longer than the
Ten new species of Lachenalia (Liliaceae) 413
lai, 17 a, Io, & Gb
4. habitat; b. two plants; c. inflorescence; d. flowers.
Lachenalia sargeantii:
414 Journal of South African Botany
outer, up to 20 mm long, 7 mm broad near the apex, slightly emarginate, cream
with a greenish-brown blotch near the apex, and a magenta keel; stamens a little
shorter than the inner segments; ovary obovoid, 3 mm diam., green; style finally
exserted. Capsule obovoid, membranous 5 mm diam; seed black, globose with a
narrow membranous decurrent aril widening toward the funicle, testa colliculate.
DIAGNOSTIC CHARACTERS
The long cream, pendulous, tubular flowers which are rounded at the base and
slightly constricted above it, are grouped at the apex of the long peduncle, on long
magenta pedicels, a character which is unusual in the genus. Its seed too is
distinctive, agreeing in shape and the colliculate testa with that of its closest ally
L. montana Schltr. ex Barker, the only two species known to have the latter
character.
Type Material: CAPE PROVINCE—3420 (Bredasdorp); (-CA), Bredasdorpberge, in
white sand among TMS boulders, 21/11/1970, W. F. Barker 10812 (NBG, holo., iso.).
Fic. 18
Lachenalia sargeantii: 1. seed side view; 2. seed from above; 3. seed from below.
This unique species has only been collected in one locality and was found
quite by accident by a young visitor from New Zealand, Robert Scott, who when
on a hitch-hiking tour of the Western Cape, decided to climb one of the mountains
near Bredasdorp in November, 1970, not long after a fire had burnt out a large part
of the area. He gathered some small portions of various plants and took them to
Mr and Mrs P. A. Sargeant, who brought them to the Compton Herbarium to be
identified. Among them were a Gladiolus sp. and an inflorescence of this
Lachenalia, which was completely new to science. Mr Sargeant very kindly
arranged a visit to the site with Robert as guide, where a collection was made of
specimens for the herbarium and some bulbs for cultivation.
They were found growing in white sand, in open spaces among TMS boulders,
on a burnt area with a northern aspect, while the Gladiolus sp. was in full flower
in crevices of large TMS rocks, where it had been protected from the fire. The
Gladiolus later proved to be one which had been collected many years before, by
Dr H. Bolus, in the same locality at the same time of year. The fact that he failed
Ten new species of Lachenalia (Liliaceae) 415
to collect the Lachenalia suggests that the area had not been burnt at the time, and
that L. sargeantii requires to be stimulated by fire before it will flower. This
supposition is strengthened by the fact that the site was visited in two subsequent
seasons, only a few scattered flowering plants were seen on the first occasion, and
none at all on the second. Up to the present time the cultivated bulbs have
produced leaves but have not flowered.
The species is named in honour of Mr and Mrs Sargeant, who are keen
mountaineers and conservationists, and have photographed the high altitude flora
of the Western Cape for many years.
SPECIMENS EXAMINED
CAPE PROVINCE—3420 (Bredasdorp): (-CA), 17/10/1970, R. Scott s.n.sub.
NBG 93894 (NBG), 21/10/1970, W. F. Barker 10802 (NBG, holo., iso.).
Lachenalia montana Schltr. ex Barker, sp. nov.
Distinguitur foliis 2 linearibus lanceolatis conduplicatis patentibus, inflorescen-
tia racemosa, pedunculo atro-marronino, pedicellis brevis patulis magenteis,
floribus cremeis campanulatis ad basim rotundatis, segmentibus externis virido-
fuscis gibbosis, segmentis internis externa paulo superantibus apice maculo
virido-fusca, staminibus paulo exsertis. seminibus globosis, arillo membranoso
undulato decurrente, testa colliculata.
Plant 100-330 mm high. Bulb globose 10—20 mm diam., with dark brown
leathery tunics produced into a short neck. Leaves 2, linear or linear-lanceolate,
conduplicate loosely clasping the base of the peduncle, suberect to spreading,
150-350 mm long and up to 20 mm broad at ground level, green tinged with
‘maroon toward apex, without markings. Peduncle 80-230 mm high, much longer
than the raceme, slender, dark maroon, glaucous or faintly blotched. Inflorescence
racemose 10 to many-flowered; flowers cernuous, concentrated into a short or long
head, sterile portion very short; bracts white or tinged with pale magenta,
lanceolate, short up to 2 mm long; pedicels very pale magenta, variable in length
2-15 mm long; in the short-headed fewer flowered inflorescences the pedicel are
usually 5 mm long, increasing markedly in length toward the apex, in the more
elongated heads the pedicels are short at the base and lengthen very gradually to
the apex. Flowers 5—9 mm long 4—5 mm diam. at the base, cernuous, campanu-
late, rounded at the base; tube 2 mm long, outer segments up to 5 mm long, ovate
cream with greenish-brown gibbosities; inner segments slightly longer than the
outer, up to 7 mm long, 4 mm broad, obovate, cream with a greenish-brown
blotch near the apex, upper edge of margin slightly recurved; buds tinged with
magenta; stamens a little exserted; ovary obovoid, green, 3 mm diam.; style up to
6 mm long, finally exserted. Capsule obovoid, membranous, 4 mm diam; seed
globose, aril membranous, undulate, decurrent, testa colliculate.
416 Journal of South African Botany
Fic. 19 a, b.
Lachenalia montana: a. habitat; b. inflorescence.
Ten new species of Lachenalia (Liliaceae) 417
DIAGNOSTIC CHARACTERS
Lachenalia montana has a number of affinities with L. sargeantii, the two
narrow conduplicate leaves loosely clasping the peduncle, the unusual capitate
type of inflorescence, with the pedicels lengthening toward the apex, and the seed
which is similar in shape and has a colliculate testa. However it differs in having
very much smaller campanulate flowers, forming a denser head, on much shorter
pedicels, and the set of the flowers is cernuous rather than pendulous.
ce
Fic. 20.
Lachenalia montana: 1. seed side view; 2. seed from above; 3. seed from below.
Type Material: CAPE PROVINCE—3419 (Caledon): Onrust, Hermanus (-CA),
16/10/1966, W. F. Barker 10485 (NBG, holo.).
The earliest known collection of this interesting species was made by Zeyher
at Houwhoek, and specimens have languished in the incertae in the collections at
Kew and at the South African Museum ever since. In November, 1896 Rudolph
Schlechter made copious collections at Houwhoek, at an altitude of 600 m, and
distributed them to many herbaria under the manuscript name Lachenalia monta-
na. Since that time a number of other collections have been made on the seaward
side of the mountains at Hangklip, Kleinmond, and Onrust near Hermanus, some
only a few feet above sea level, so that the name is not altogether apt, but it seems
expedient, as it has been so widely distributed under the name, to retain it.
Many of the recent collections were made on burnt ground, and at Onrust
where it grew and flowered in great profusion after a fire in October 1966, it was
observed to decrease in flowering capacity in subsequent seasons, as other
vegetation recovered and became dominant.
SPECIMENS EXAMINED
CAPE PROVINCE—3319 (Worcester): Louwshoek Peak (-CC), 15/12/1944,
E. Esterhuysen s.n.sub (BOL).
3418 (Simonstown): Hangklip (-BD), 6/12/1950, J. S. Linley s.n.sub. SAM
61011 (SAM); 6/12/1950, Werner s.n.sub. NBG 72391 (NBG).
——3419 (Caledon): Onrust (-AC), 30/10/1962, Thomas s.n.sub. NBG 70038
(NBG); Onrust near Hermanus, 16/10/1966, W. F. Barker 10485 (NBG, holo.);
418 Journal of South African Botany
Fern Kloof Hermanus (-AD), 11/1/1972 (fruiting), T. T. Barnard (NBG); Klein-
mond, 30/11/1962, Cloete s.n.sub. NBG 70035 (NBG); Houwhoek (-BA), De-
cember, Zeyher 4294 (SAM, K); Houwhoek, 24/9/1896, R. Schlechter 9403
(BOL, BM, G, GRA, K, L, PRE, S, Z).
ACKNOWLEDGEMENTS
In the preparation of this paper I am indebted particularly to Professor E. A.
Schelpe who has made the facilities at the Bolus Herbarium available to me since
my retirement in 1972, and for his advice and assistance in many other ways,
including the preparation of the Latin translations. My sincere thanks also to
Professor R. H. Compton for his assistance with the Latin descriptions of the
seeds. Dr J. P. Rourke has been most helpful in making my research material in
the Compton Herbarium available to me, and in his advice on matters concerned
with publication. Mrs. C. Labuschagne kindly prepared the Afrikaans abstract.
I wish to express my thanks to all curators of the herbaria mentioned, who
have provided me with facilities or sent specimens on loan, to the Photographic
Department of the Jagger Library for the photographs of the paintings, and to the
British Museum (Nat. Hist.) and the Bolus Herbarium for the use of the
photographs of their herbarium sheets. My grateful thanks to all collectors
particularly Miss E. Esterhuysen, Mrs M. Thomas, Mr F. Stayner and Mr and Mrs
P. A. Sargeant.
Jl S. Afr. Bot. 44 (4): 419-420 (1978)
THE IDENTITY OF POLYPODIUM POPPIGIANUM METT. (F ILICES)
J. P. ROURKE
(Compton Herbarium, Kirstenbosch)
E. A. C. L. E. SCHELPE
(Bolus Herbarium, University of Cape Town)
ABSTRACT
__ The name Polypodium péoppigianum Mett. is considered applicable to a small gram-
mitid fern from Victoria Peak, Cape Province, Grammitis poeppigana (Mett.) Schelpe
comb. nov. is proposed. In the absence of the type, Esterhuysen 29290 is proposed as a
neotype.
UITTREKSEL
DIE IDENTITEIT VAN POLYPODIUM POPPIGIANUM METT. (FILICES)
Die naam Polypodium péppigianum Mett. word as toepaslik vir ’n klein grammitiede
varing vanaf Victoriapiek, Kaapprovinsie beskou. Grammitis poeppigana (Mett.) Schelpe
comb. nov. word voorgestel. Omdat ’n tipe ontbreek word Esterhuysen 29290 as neotipe
voorgestel.
In 1875, Mettenius, in his treatment of the genus Polypodium, as then
construed, described a Polypodium pdppigianum as follows:
“‘Rhizoma adscendens, caespitosum, paleis lanceolatis, acuminatis onustum;
folia 1-1%2"’ ‘longa, 2%4’’’ lata coriacea, glabra, spathulato-lanceolata, obtusa;
nervi immersi, sori ad basin rami antici, breviter oblongi, costae subparalleli,
denique costam ipsam tegentes.
Prom.b.spei (in fissuris rupium in fauce umbrosa montium Hott. holland.)
Poppig.””
In 1943, Prof. C. L. Wicht of Stellenbosch discovered a small grammitid fern
on Victoria Peak and this was referred by Alston & Schelpe (1952) to Polypodium
magellanicum (Desv.) Copel. It was only realised later that this could represent the
‘‘lost’’ P. péppigianum Mett., but Poppig was not known to have collected in
South Africa. However, the clue was provided in a letter from Poppig in the
Hooker correspondence at Kew, that Poppig maintained a collector at the Cape.
This collector has now been identified as Wilhelm Gueinzius who collected at the
Cape between 1839 and 1841. Furthermore, he collected in the Hottentots Holland
Mountains, where he discovered Leucospermum gueinzii Meisn. and could have
ascended Victoria Peak, the highest peak in the vicinity.
Miss E. Esterhuysen, who has re-collected this fern on Victoria Peak, records
that it only grows in horizontal crevices on the walls of a south aspect gully in
Accepted for publication 23rd August, 1978.
419
420 Journal of South African Botany
cliffs at altitudes between 1 450 and 1 500 m. This agrees remarkably well with
the habitat description given by Mettenius.
No other grammitid fern has been found in South Africa and little doubt
remains that the name P. poéppigianum refers to the plant found on Victoria Peak.
Unfortunately the holotype is not in Vienna with Poppig’s herbarium, and has not
been traced in any other European herbaria. It is presumed lost, or destroyed
during World War II.
This south-western Cape fern bears the earliest specific epithet applicable to a
widespread Southern Hemisphere fern (Parris, 1975) with an intricate synonomy.
Consequently a new combination, Grammitis poeppigana (Mett.) Schelpe is
proposed (basionym-Polypodium poppigianum Mett., Farngatt., Polypol.: 37, No.
17 (1875)). In the absence of a holotype, Esterhuysen 29290 (BOL) from Victoria
Peak is proposed as a neotype. Isoneotypes have been or will be placed in Kew,
Edinburgh and St. Louis.
REFERENCES
Aston, A. H. G. and SCHELPE, E. A. C. L. E., 1925. An annotated check-list of the
Pteridophyta of Southern Africa. Jl S.Afr. Bot. 18: 153-176.
Parris, B. S., 1975. A revision of the genus Grammitis Sw. (Filicales: Grammitidaceae) in
Australia. J. Linn. Soc. Bot. 70; 21-43.
JIS. Afr. Bot. 44 (4): 421 (1978)
Book REVIEW
MESQUITE: ITs BIOLOGY IN Two DESERT ScRUB ECOsysTEMs, edited by B. B.
Simpson, with pp. 250. New York, Dowden, Hutchinson & Ross, Inc., 1977.
$22,00.
Scientific investigation into the problems concerning our veld takes rather a back seat
in South Africa. Veld research often entails long-term projects which are avoided by the
average researcher and student.
Is the presence of the mesquite (Prosopis spp.) an asset to the arid veld types of South
Africa and South West Africa or can it be considered an introduced pest and a danger to the
carrying capacity of the veld? It was introduced into Southern Africa many years ago but
one hears very little about it in the conference room or in literature.
Mesquite: Its Biology in Two Desert Scrub Ecosystems is a very welcome American
contribution to the literature on veld ecology that we in South Africa can also make good
use of. The book covers tremendous ground in research by teams of researchers. Topics
studied and reported on include: Phenology, morphology and physiology of mesquite:
Prosopis leaves, flowers and fruits as a resource for insects and invertebrates; Its food,
medicinal cosmetic, recreational, fuel and other values; The spread of mesquite; Factors
involved in its spread; Control of mesquite. There is also an annoted key to the species of
the world of the genus Prosopis.
We in South Africa can probably not afford research on as large a scale as the
Americans and must therefore make good use of this very comprehensive publication.
The text is adequately supported with illustrations, graphs, maps and photographs. It is
2 neat, wellbound book comprising 250 pages and measuring 240 x 160 mm. The reader is
supplied with a very comprehensive and impressive reference list. The book has an
adequate subject index as well as an index of scientific names. It will appeal to the
undergraduate and the post-graduate student as well as the research worker.
May this book help to stimulate more interest in our South African veld and ecology
and the influence of modern man on its destiny.
J. G. V. JOUBERT
42]
JOURNAL OF SOUTH AFRICAN BOTANY
VOLUME 44
1978
INDEX OF PLANT NAMES
Note: Plant names appearing in tables or lists are not included in this index. Page references to new
taxa are printed in bold type. An asterisk indicates an illustration.
PAGE
Acmadenia candida Williams .......... 352
Agkiwanensisawilliamss esac eeeenie 347
/\. TANGENTS, Gocaooeoepocenonoes 350
A. patenti folia Williams .......... 343, *346
A. wittebergensis (Compton) Williams .. 359
Asplenium erectum Bory ex Willd. .163, *165
Athyrium scandicinum (Willd.) Presh. *180, 184
Brabeium stellatifolium L. ............ 143
Cassia italica (Mill.) Lam. ex F. W. Andr. 67
C. italica (Mill.) Lam. ex F. W. Andr.
ssp. arachoides (Burch.) Brenan ...... *68,
1D), 7A, 7/7)
C. italica (Mill.) Lam. ex F. W. Andr.
ssp. micrantha Brenan....... *68, 4705 4711
Diastella buekii (Gandoger) Rourke ..... 58
Dryopteris inaequalis (Schlechtend.)
Kt Zen eie rac haisicransierer ernie: 157, *158
Encephalartos caffer Mig. ............. 84
E. eugene-maraisii Verdoorm....... S175 +3119
E. lehmannii Ecklon ex Lehm.......... 84
Enrica junonia Bolus). =: -2--...-.----.- 97
Euchaetis avisylvana Williams 339, *342, *343
Earintonsaawillianisicyyereei rere 335, *338
JB. IEE TTA, cconcanccenpo00000 333
E. meridionalis Williams.............. 329
Galiiinilk, co concopacoostegnnoatdoeu so 219
G. amatymbicum Eckl. & Zeyh. ....... 245
Gebredasdorpense/ butter see ea 244
G. bussei K. Schum. & K. Krause...... 246
G. bussei K. Schum. & K. Krause var.
LIKES «cososnpooupuoecenonduaddo® 247
G. bussei K. Schum. & K. Krause var.
glabrum Brenan ............-.-+--- 247
Gcapense Thunb) =. -2.-------------- 228
G. capense Thunb. ssp. capense ... .*211, 230
G. capense Thunb. ssp. garipense (Sond.)
Wiittar oor odocamob eau dTUapOO.OnS 236
G. capense Thunb. ssp. garipense (Sond.)
Puff var. garipense ........-..-+--- 236
G. capense Thunb. ssp. garipense var.
wittbergense (Sond.) Puff ..........- 237
G. capense Thunb. ssp. namaquense (Eckl.
ie TESS Nig coococenastoc0ue *212, 234
G. chloroionanthum K. Schum...... *217, 269
G. monticolum Sond. .........------- 243
G. mucroniferum Sond.........-..---- 259
423
PAGE
Galium mucroniferum Sond. var. dregeanum
(SondS)iRutianee eee eee 260
G. mucroniferum Sond. var. muc-
RMGUAN sooronrcaccnooerocancacr 259
G. scabrelloides Puff .. .*212, *215, #218, 250
G¥scabrellumpke;Schummesoee seer
~G. spurium L. ssp. africanum Verdc. .... Dl,
GysubvillosumySond sero eae IIB), PSp)
G. subvillosum Sond. var. subglabrum
Putfiycsytevioiets Geasiacen Secs ce-sale eee
subvillosum Sond. var. subvillo-
SUIM Wee ycus, ot on -eesron een ReT PINNAD, ADI}, 72)I/
G. thunbergianum Eckl. & Zeyh. .. .*217, 221
G.
G. thunbergianum Eckl. & Zeyh. var.
hirsutume(Sonds) Verdes eae 2)
G. thunbergianum Eckl. & Zeyh. var.
thunbergianunieeeece eet ieee: 222
G. tomentosum Thunb. .*211, *215, 263, *266
GeundulatumiBuiteeneeeece ecco: 262
Grammitis poeppigana (Mett.) Schelpe .. 419
Jubaeopsis caffra Becc. ........... 127, *129
Kalanchoe luciae Hamet ssp. luciae ..... 89
K. luciae Hamet ssp. montana (Compt.)
TONE cooccaoavensosbeangeoono” 89
Kemeglectawllioelkenieaeet reer 90
Kesrubineaslhocikenterrerer erecta irr 90
Lachenalia concordiana Schltr. ex Barker 407,
*408
leacongestayBarkenamrecraerr irr 399, *400
L. esterhuysenae Barker........... *397, 398
L. glaucophylla Barker ...... 409, *410, *411
L. montana Schltr. ex Barker ...... 415, *416
WW. multifolia Barker. + -¢4: 25-1 394, *395
L. namaquensis Schltr. ex Barker . . 402, *403
L. peersii Marl. ex Barker......... 391, *392
I, sargeantii) Barket)......-------1-- 412, *413
IL, (ip eoltore (UL,)) WWD), cogcooovcecgnon 111
Ie) verticillata’ Barker) sneer ee: 405, *406
Leucadendron flexuosum Williams..... . 58
IL, sikorataliton RUB coccsonudns0ncGa000% 58
L. levisanus (L.) Berg. ...........---- 58
L. verticillatum (Thunb.) Meisner ...... 58
lv. xanthoconus ON Ktze err 143
Lycopodium gnidioides L.f. ....... 157, *158
Macrostylis ramulosa Williams .... - 356, *359
424 Index to Plant Names
PAGE
Papillaria natalensis Sim.............. 300
Pectinaria exasperata Bruyns....... 153, #155
Refreanvolubilisviussseem ere 119, 120, 122
Polypodium poppigianum Mett. ........ 419
Polystichum luctuosum (Kunze) Moore 166,
*167
12% mec (ES) IDIGG soceosc00n0000 166
12, UINTINS GIVING cocoaccccccns 177, *180
P. setiferum var. fuscopaleaceum sensu
Schelpeyetc heresies erie: 73, 7S)
P. transkeiense Jacobsen .......... 169, *171
P. zambesiacum Schelpe .......... 167, *171
Protea inopina Rourke................ 376
I EATON TWN) occocavccdo0000n06 143
P. lepidocarpodendron L. ............. 143
PAGE
Protea longiflora Lam. ssp. potbergensis
Rourkes's |, acc ae aerent cys ie eer 378
1, MUCO MO SANS, cococovccege0c0c 363
nubipenaROUTKe sere eeerner 373
Paodoratay@i hun Sees eee ere 363
12, OOSMA IROWUME soccocsccge00000005 375
P. roupelliae Meisn. ssp. hamiltonii Beard
ex Rourkekcnsnes nate Se 378
NetruniarciliatayReB laeee eee eee 58
Sb GYeMOlES IRIE soocsedacavog00000c 363
Sb 1WMESIRA RIT, coscoccsgccocccc006 363
S. linearis Salisb. ex Knight ........... 363
Sesclophal kai ghtlaeenen eee eee 363
Strelitziagreginaey Altes 103
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